George Srzednicki (Editor) - Chaleeda Borompichaichartkul (Editor) - Konjac Glucomannan-Production, Processing, and Functional Applications-CRC Press (2020)

Konjac Glucomannan
Konjac Glucomannan
Production, Processing,
and Functional Applications
Edited by
George Srzednicki
Chaleeda Borompichaichartkul
CRC Press
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Library of Congress Cataloging‑in‑Publication Data
Names: Srzednicki, G. (George), editor. | Borompichaichartkul, Chaleeda,

editor.
Title: Konjac glucomannan : production, processing, and functional
applications / edited by George Srzednicki, Chaleeda
Borompichaichartkul.
Description: Boca Raton : CRC Press, [2020] | Includes bibliographical
references and index.
Identifiers: LCCN 2019060021 (print) | LCCN 2019060022 (ebook) | ISBN
9781138367173 (hardback) | ISBN 9780429429927 (ebook)
Subjects: LCSH: Polysaccharides. | Polysaccharides industry. | Konjak. |
Amorphophallus.
Classification: LCC TP248.65.P64 K66 2020 (print) | LCC TP248.65.P64
(ebook) | DDC 664/.06--dc23
LC record available at https://lccn.loc.gov/2019060021
LC ebook record available at https://lccn.loc.gov/2019060022
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This book is dedicated to the memory of late Dr. James F. Maxwell (‘’Max’’), one of the most
prominent botanists of Thailand and neighbouring countries. Max, a native of New York State,
graduated from Ohio State University with a BSc in botany and later with an MSc from the
University of Singapore, spent over 40 years in Thailand where he worked as a taxonomist in
various universities. I came to know Max during the first Flora Malesiana symposium, which
held in Leiden 1989. Our post-conference meeting lasted a brief 20 minutes after which we
continued our conversations through letters, faxes and emails for almost the next 30 years and
never came face-to-face again. He promised to look out for Amorphophallus species during his
field trips and so provided a hugely important contribution to my taxonomic work on this genus
over the years. I was pleased to honour him in naming the most spectacular species in Thailand
after him, Amorphophallus maxwellii Hett., a discovery by himself. Later he co-authored
with me the new species named as A. hemicryptus Hett. & Maxw found in a small island
in the Mekong on Cambodian soil. As Max was always in need of photocopies from botanical
literature and CDs with mostly female opera singers of long forgotten times, there was a lively
exchange of such objects and he sent Amorphophallus tubers for my research collection.
His thousands of herbarium specimens collected during his field trips in Thailand, Laos,
Cambodia and Myanmar are spread across herbaria in the world, many in the Chiang Mai
University herbarium where he held office as curator and from where he roamed the tropical
forests. His knowledge of plant ecology in those forests was phenomenal. As a mutual
acquaintance, Martin van den Bult (pers. Comm.), joined him in a number of field trips and told
me how the high standards of working Max held, which often led to outcries after seeing lesser
qualitative work, using such terms as “moron” and “cretin” but always in good spirit (because
Max liked to drink a strong local “schnapps”, which he called “Mekong”). His relationship
with several Thai botanists was brittle, to say the least but in the long run, many of them
and an equally high number of students gained profuse knowledge in the field of botany.
I deeply cherish my giant pile of faxes and emails from Max in all their good
and sometimes evil spirit. A unique person suddenly left us when Max died
on 12 May 2015 of a massive heart attack during forest work. I imagine that
dying during field work could have been something he would wish for.
A fine obituary has been published in Biotropica 48(1) in 2016.
Wilbert Hetterscheid
One of the last photographs of Max during his field survey in Rayong Province.
Max and Amorphophallus maxwellii

Contents
Preface...............................................................................................................................................ix
Editors...............................................................................................................................................xi
Contributors.................................................................................................................................. xiii
1. Introduction..............................................................................................................................1
Chaleeda Borompichaichartkul and George Srzednicki
2. Botanical Background to Amorphophallus........................................................................ 5

Wilbert Hetterscheid, Li Heng, Wang Zhonglang, Orachorn Mekkerdchoo,
and Cyrille Claudel
3. Biosynthesis and Decomposition of Konjac Glucomannan....................................... 101

Orachorn Mekkerdchoo, Yu Lei, and Zhao Jianrong
4. Field Production of Konjac................................................................................................ 115

Zhang Shenglin, Jiang Xuekuan, and Hadi K. Purwadaria
5. Postharvest Technology of Konjac................................................................................... 161

Rarisara Impaprasert, Zhao Jianrong, George Srzednicki, Yu Lei, and Tao Ruixuan
6. Processing of Konjac Flour................................................................................................ 173

Rarisara Impaprasert, Zhao Jianrong, and George Srzednicki
7. Physico-Chemical Properties of Konjac Glucomannan.............................................. 189

Patricia Le Bail, Céline Lafarge, and Nathalie Cayot
8. Advances in Drying Technology...................................................................................... 209

Lamul Wiset, Nattapol Poomsa-ad, Rarisara Impaprasert,
and Chaleeda Borompichaichartkul
9. Konjac Industry in Major Producing Countries........................................................... 223

Zhang Shenglin, Hadi K. Purwadaria, Chaleeda Borompichaichartkul,
and Phattanit Tripetch
10. Applications of Konjac Glucomannan in Food and Medicine................................... 255

Chaleeda Borompichaichartkul, Afwa Hayuningtyas, and Phattanit Tripetch
viii Contents
11. New Trends in the Konjac Flour Industry...................................................................... 265

Chaleeda Borompichaichartkul, Desi Sakawulan, Patthasarun Pruksarojanakul,
12. Concluding Remarks.......................................................................................................... 277

Index.............................................................................................................................................. 279
Preface
There are about 200 species of genus Amorphophallus in the Araceae family. Except for
about 25 species native to Africa, most of them originate from southern, south-eastern and
eastern Asia, northern Australia, or Pacific islands. The underground tubers of several of
the Asian species are rich in glucomannan, which is a water-soluble polysaccharide that
is considered as a dietary fibre. As such, it resists human digestive enzymes and absorp-
tion in the small intestine. Hence, it is used as food or food additives (e.g., thickener or
emulsifier), food supplements for weight management, and also in the pharmaceutical
and cosmetic industries. Given these properties, it is an important economic raw material.
The natural sources of glucomannan besides Amorphophallus include tubers of orchids of
genus Orchis, hemicellulose present in the wood of conifers, and some bacterial and yeast
cell walls. However, the by far most important commercial source of glucomannan are vari-
ous species of the genus Amorphophallus. One of the main species producing glucomannan
is Amorphophallus konjac, from which the common name of this crop ‘konjac’ originates.
The generic name of this compound is ‘konjac glucomannan’, generally designated by the
acronym KGM. China had owned and utilized abundant glucomannan resources as early
as 2000 years ago, but until the mid-1980s, the konjac production was limited to plants
growing in the wild or planted in gardens in front of the houses. After 1985, with the
advancement of science and technology and the development of foreign trade, responding
to the increasing konjac flour demand in the international market, people began to recog-
nize the potential value of konjac-derived products, and the development and utilization
of konjac resources took rapidly off. After nearly 30 years of relentless efforts by konjac
researchers, growers, and processors in the konjac industry, China has formed a complete
konjac supply chain, and its industrial scale continues to expand. In 2009, konjac flour
production from China exceeded that of Japan’s which until then was the number one
producer in the world.
Japan was the first country to form a complete konjac supply chain. About 1500 years
ago, konjac was introduced to Japan from China by Buddhist monks. Although its cultiva-
tion and utilization took place later than in China, the konjac industry focusing mainly on
the production of jelly-like cake called konnyaku, started in Japan earlier than in China.
In Japan, konjac has been traded as a commodity on the market since the fourteenth
century. The konjac cultivation evolved from a natural perennial growth to the annual
cultivation after the seventeenth century. Since then, konjac has been considered as a
food source of national importance, and its cultivation and processing became encour-
aged since the nineteenth century. Konjac became one of the main ingredients in Japanese
healthy traditional meals in the 1960s. The konjac plantation areas ranked first in the world
by the 1970s. By the 1990s, the output of fresh harvested tubers reached 100,000 tonnes and
12,000 tonnes of konjac flour was produced.
Although the centre of origin and the production areas of several glucomannan producing
species are in Southeast Asian countries, the utilization of konjac is mainly driven by the
demand of Chinese and Japanese markets. In the past, it was mainly used to make k onjac
cake by harvesting the subterranean tubers, called ‘corms’, in their natural habitat, sun
drying them after slicing, and then exporting them to the Chinese market. In the last decade,
because of the increased investment in processing equipment from China and Japan, the
konjac cultivation and mechanical drying of chips has increased in Southeast Asia.
ix
x Preface
Konjac is a very particular plant. It has originated in the lower layers of tropical forests
in Asia, and thus has the characteristics of plants growing in warm and moist environ-
ments, half shade and does not support neither dryness nor dampness. Underground
tubers are particularly susceptible to bacterial soft rot disease (Erwinia carotovora) infection
that causes devastating damage. Konjac requires special processing equipment because
the underground tubers have a high moisture content and konjac glucomannan is very
prone to browning.
The high glucomannan content of konjac underground tubers has attracted wide attention
by the international community. The United States opened its market in 1997, giving the
konjac flour the status of ‘generally recognised as safe’. The EU recognized konjac flour as a
food additive in the EU in 1998 and 2003. This development stimulated the research on the
konjac applications in the European Union. Subsequently, there is a growing demand for
konjac glucomannan in the global market.
The konjac industry has gradually developed thanks to the continuous scientific and
industrial research in China and Japan for nearly 30 years. Although the potential benefits
of konjac glucomannan for the health of people in modern society are considerable, the
popularity of konjac is still below that of most of the other major crops. Therefore, in order
to fill this gap, it is necessary to summarize and update the information on the current
status of konjac field production, the processing industry, and applications in food and
other industries.
The authors of this book are researchers and experts who have been engaged in the research
and development of the konjac industry for many years. This book includes the results of
laboratory, field, and industrial research in the konjac producing and consuming countries.
It includes the latest information on the biology of the Amorphophallus species producing
glucomannan, agronomy, raw materials processing equipment, the applications in food
and other industries, and the related processing technology. It describes the most advanced
cultivation and processing technologies of the konjac industry in various countries. It is
expected that the information covered by this book will be useful as a reference for the sci-
entists and konjac industry practitioners. However, because of our limited ability to obtain
all the relevant data, especially from the industrial sources, we would like to apologize if
there are still gaps in the information provided.
Prof. Zhang Shenglin

Chinese Konjac Research Institute, Chongqing
Editors
Dr George Srzednicki completed a PhD in technical sciences

with major emphasis on agricultural engineering in 1977
at the Swiss Federal Institute of Technology (ETH, Zurich,
Switzerland). He started his professional career as a consul-
tant for engineering companies (Elektrowatt, Agroprogress)
in Africa (Senegal, Nigeria), and Asia (Bangladesh), and
also for the Swiss government agency funding technical
cooperation projects in Central America (Costa Rica). His
activities included harvest and postharvest techniques with
a focus on the storage of food grain, seed, and perishable
crops. In 1988, he joined the University of New South Wales
(Department of Food Science & Technology, later School of
Chemical Engineering) in Sydney (Australia). He became
involved in a range of research projects such as design of food processing equipment,
especially drying, process control, and optimization, product quality in food dehydra-
tion processes, energy efficiency in food processing, etc. Several research projects were
conducted in collaboration with research organizations in countries such as Thailand,
the Philippines, Malaysia, Vietnam, Indonesia, China, India, and Papua New Guinea.
He supervised 24 PhD theses (five related to konjac glucomannan), was involved in the
development of training materials and their delivery, authored 4 book chapters, and over
130 papers for peer-reviewed journals and conference proceedings.
Assoc. Prof. Dr Chaleeda Borompichaichartkul completed
a PhD in food science and technology with major emphasis
on food processing and engineering especially in drying tech-
nology in 2004 (University of New South Wales, Australia).
In 2004, she started to work at the Department of Food
Technology, Faculty of Science, Chulalongkorn University in
Bangkok as a junior lecturer, and then, in 2005, she obtained
a postdoc scholarship from UMR-GENIAL Unité Mixte de
Recherche Genial Cemagref, ENSIA, INAPG, INRA, Massey,
France. In 2006, she created a research team on konjac glu-
comannan processing and applications at the Department
of Food Technology, Faculty of Science, Chulalongkorn
University. Her main research interests include applied
research in drying technology and extension focusing on konjac glucomannan, its process-
ing, and applications in functional ingredients and foods. The research activities deal with
studies of the physical and chemical properties of konjac glucomannan, microencapsulation
process, and development of functional films using konjac glucomannan as a raw material.
She has directed and evaluated Thai and international research projects, supervised doctoral
theses, trained national and international students, and collaborated with research groups in
Australia, New Zealand, Italy, Austria, Germany, the USA, China, Vietnam, Indonesia, and
Malaysia. To date, she has published over 40 peer-reviewed research articles (H-index of 8).
xi
Contributors
Chaleeda Borompichaichartkul Zhao Jianrong

Faculty of Science College of Agronomy
Department of Food Technology Urban Modern Agriculture Engineering
Chulalongkorn University Research Center
Bangkok, Thailand Kunming University
Kunming, China
Nathalie Cayot
Centre des Sciences du Goût et de Céline Lafarge
l’Alimentation Centre des Sciences du Goût et de
Université de Bourgogne l’Alimentation
Dijon, France Université de Bourgogne
Dijon, France
Cyrille Claudel
Department of Biology Patricia Le Bail
Institute for Plant Science and Unité de Physicochimie des
Microbiology Macromolécules
University of Hamburg Institut National de la Recherche
Hamburg, Germany Agronomique
Nantes, France
Afwa Hayuningtyas
Faculty of Science Yu Lei
Department of Food Technology College of Agronomy
Chulalongkorn University Urban Modern Agriculture Engineering
Bangkok, Thailand Research Center
Kunming University
Li Heng Kunming, China
Kunming Institute of Botany
Chinese Academy of Sciences Orachorn Mekkerdchoo
Kunming, China Faculty of Agro-Industry
King Mongkut’s Institute of Technology
Wilbert Hetterscheid Ladkrabang
Von Gimborn Arboretum Bangkok, Thailand
Doorn, the Netherlands
Nattapol Poomsa-ad
Rarisara Impaprasert Faculty of Engineering
Faculty of Science Mahasarakham University
Department of Microbiology Mahasarakham, Thailand
King Mongkut’s University of Technology
Thonburi
Bangkok, Thailand
xiii
xiv Contributors
Patthasarun Pruksarojanakul George Srzednicki

Department of Food Technology School of Chemical Engineering
Faculty of Science The University of New South Wales
Chulalongkorn University Sydney, Australia
Bangkok, Thailand
Phattanit Tripetch
Hadi K. Purwadaria Faculty of Science
Departemen Teknik Mesin dan Biosistem Department of Food Technology
Fakultas Teknologi Pertanian Chulalongkorn University
Kampus Institut Pertanian Bogor Bangkok, Thailand
Bogor, Indonesia
Lamul Wiset
Tao Ruixuan Faculty of Engineering
School of Foreign Languages Mahasarakham University
Yunnan Minzu University Mahasarakham, Thailand
Kunming, China
Jiang Xuekuan
Desi Sakawulan The Konjac Association of Chinese Society
Faculty of Science for Horticultural Science
Department of Food Technology Chongqing City, China
Chulalongkorn University
Bangkok, Thailand Wang Zhonglang
Kunming Institute of Botany
Zhang Shenglin Chinese Academy of Sciences
College of Horticulture and Landscape Kunming, China
Architecture of Southwest University
(SWU)
The Konjac Association of Chinese Society
for Horticultural Science
Chongqing City, China
1
Introduction
Konjac glucomannan (KGM) is a high-molecular-weight polysaccharide that was originally

extracted from the corms (underground storage organs) of Amorphophallus konjac K. Koch,
a perennial plant belonging to the family Araceae. Glucomannan consists of D-mannose
and D-glucose units at a molar ratio of 1.6:1.0, connected by (1-4)-glycosidic bonds and is
a water-soluble hydrocolloid obtained from konjac flour. Konjac flour is the unpurified
raw product from the corms of various species of Amorphophallus as per specifications
for konjac flour [INS 425] of the Joint FAO/WHO Expert Committee on Food Additives.
Konjac flour is also known under the synonyms konjac mannan, konjac, konnyaku, and
konjac glucomannan (JECFA 2006). The European Union authorizes the use of purified
forms of glucomannan, namely, konjac gum (E 425 i) and konjac glucomannan (E 425 ii) as
food additives; konjac gum is obtained by aqueous extraction, while konjac glucomannan
is obtained by washing with water-containing ethanol (EFSA 2017). The chemical charac-
teristics of glucomannan are described in the Merck Index (2006) and in the CAS registry
under the number 37220-17-0 (EFSA 2017).
Besides the genus Amorphophallus, there are also other sources of glucomannan, such
as the roots of certain orchids, cell walls of mung bean seedlings (Elbein 1969), bacteria,
yeasts (Chorvatovičová et al. 1999; Tokoh et al. 2002), and also the hemicellulose in the
wood of some conifers and angiosperm trees. However, given their high glucomannan
content and the relatively simple extraction technique, the main commercial sources of
glucomannan are the various species of Amorphophallus.
In the last few years, KGM drew the particular attention of scientists and the food industry
due to its bioactive properties, biodegradability, and hydrophilic ability. Multidirectional
research is focusing on KGM and its derivatives in fields such as food science and nutri-
tion, biotechnology, pharmacology, and fine chemicals (Behera & Ray 2016).
As far as food science and nutritional aspects are concerned, the main areas of research
are anti-oxidant and prebiotic activity, functional foods, food additives, and their deriva-
tives. With regard to biomedical research, anti-obesity therapy, regulation in lipid metabo-
lism, laxative effect, anti-diabetic, anti-inflammatory, prebiotic, and also wound dressing
applications should be mentioned (Behera & Ray 2016).
Various species of the genus Amorphophallus have been used over the centuries in trop-
ical and subtropical Asia as a food source and as a traditional medicine. Konjac was first
described as a medicinal herb in the Shen Nong Materia Medica during the Western Han
Dynasty (206 BC–08 AD) in China. In general, corms were washed, peeled, sliced, dried, and
ground to produce konjac flour. Konjac flour was consumed in the form of cake (or gel) after
boiling the flour with plant ash. In traditional cuisines, konjac was also consumed in the form
of noodles, tofu, and snacks or as konjac curd, which was usually braised with meat. In tra-
ditional Chinese medicine, konjac gel has been used for the treatment of asthma, coughs,
hernia, breast pain, burns, as well as haematological and skin disorders (Chua et al. 2010).
1
2 Konjac Glucomannan
In the sixth century AD, konjac was introduced into Japan and Korea, initially, as a
medicinal plant. Later on, konjac became popular as a vegetarian foodstuff. For culinary
use, konjac flour is pounded with lime and water into a gelatinous grey cake, a key ingredi-
ent in Japanese noodles (shirataki) and cuisines, such as sukiyaki and gyudon (Chua et al.
2010). More recently, under the influence of the konjac industry in Japan and China after
World War II, the production of glucomannan flour expanded to Southeast Asia, particu-
larly, Thailand and Indonesia (Impaprasert et al. 2014; Yanuriati et al. 2017). The current
world konjac flour production is in excess of 25,000 tons (Parry 2010). The largest producers
of konjac flour are China and Japan, these two countries account for approximately 60%
and 28% of global production, respectively (Liu 2004).
In spite of the wide use of KGM as a foodstuff and in traditional medicine in the Orient,
its consumption in the West is relatively limited. This is due to the fact that knowledge
about the properties and potential applications of KGM and its derivatives is still rather
limited compared to other well-established polysaccharides, such as cellulose, starch, and
chitosan. However, since the United States Food and Drug Administration listed konjac
flour as generally recognised as safe (GRAS) (FDA 1997), shortly followed thereafter by
recognition as a food additive in the European Union (EU 1998; Parry 2010; EFSA 2017),
KGM became increasingly used in emulsifier and stabilizer products for the food, drinks,
cosmetic, and pharmaceutical industries. Therefore, the aim of this book is to provide an
overview of the botanical background of the glucomannan producing Amorphophallus spe-
cies, the biosynthesis of glucomannan, field production, processing techniques, status of
KGM processing industry in the main producing countries, application of KGM-derived
products, and the current research leading to the development of new products. It is
expected that this book will be a useful reference for researchers, members of the industry
including growers and processors, and also all potential users of products derived from
this multifunctional natural polymer.
References
Behera, S. S., & Ray, R. C. (2016). Konjac glucomannan, a promising polysaccharide of Amorphophallus
konjac K. Koch in health care. International Journal of Biological Macromolecules, 92, 942–956.
Chorvatovičová, D., Machová, E., Šandula, J., & Kogan, G. (1999). Protective effect of the yeast
glucomannan against cyclophosphamide-induced mutagenicity. Mutation Research/Genetic
Toxicology and Environmental Mutagenesis, 444(1), 117–122.
Chua, M., Baldwin, T. C., Hocking, T. J., & Chan, K. (2010). Traditional uses and potential health
benefits of Amorphophallus konjac K. Koch ex N.E.Br. Journal of Ethnopharmacology, 128, 268–278.
Elbein, A. D. (1969). Biosynthesis of a cell wall glucomannan in mung bean seedlings. Journal of
Biological Chemistry, 244(6), 1608–1616.
EFSA [European Food Safety Authority]. (2017). Re-evaluation of konjac gum (E 425 i) and konjac
glucomannan (E 425 ii) as food additives. EFSA Journal, 15(6), 1–43.
EU. (1998). Directive 98/72/EC published 4th Nov 1998 modifying European Parliament and Council
Directive 95/2/EC of 20 February 1995 on food additives other than colours and sweeteners.
FDA. (1997). GRAS title 21 CFR 170.30
Impaprasert, R., Borompichaichartkul, C., & Srzednicki, G. (2014). A new drying approach to enhance
quality of konjac glucomannan extracted from Amorphophallus muelleri. Drying Technology: An
International Journal, 32(7), 851–860.
Introduction 3
JECFA (Joint FAO/WHO Expert Committee on Food Additives). (2006). Konjac flour. Combined
Compendium of Food Additive Specifications: All Specifications Monographs from the 1st to the 65th
Meeting (1956–2005). Rome, Italy: Food and agriculture organization of the United Nations.
Liu, P. Y. (2004). Konjac. Beijing, China: China Agricultural Press (in Chinese).
Merck Index. (2006). Konjac Gum and Konjac Glucomannan. 14th ed., Merck & Co., Whitehouse Station,
NJ.
Parry, J. (2010). Konjac glucomannan. In: Imeson, A. (ed.) Food Stabilisers, Thickeners and Gelling
Agents. Singapore: Blackwell Publishing, pp. 198–215.
Tokoh, C., Takabe, K., Sugiyama, J., & Fujita, M. (2002). CP/MAS 13C NMR and electron diffraction
study of bacterial cellulose structure affected by cell wall polysaccharides. Cellulose, 9(3–4),
351–360.
Yanuriati, A., Marseno, D. W., Rochmadi, R., & Harmayani. E. (2017). Characteristics of glucoman-
nan isolated from fresh tuber of Porang (Amorphophallus muelleri Blume). Carbohydrate Polymers,
156, 56–63.
2
Botanical Background to Amorphophallus
Wilbert Hetterscheid, Li Heng, Wang Zhonglang, Orachorn Mekkerdchoo,

and Cyrille Claudel
CONTENTS
2.1 Evolutionary and Genetic Aspects.......................................................................................6
2.2 Taxonomy.................................................................................................................................9
2.2.1 Amorphophallus: Genus Description..................................................................... 11
2.3 Glucomannan-Producing Species...................................................................................... 14
2.3.1 Amorphophallus albus Liu & Wei............................................................................ 14
2.3.2 Amorphophallus amygdaloides Hett. & Sizemore.................................................. 16
2.3.3 Amorphophallus asterostigmatus Bogner & Hett................................................... 19
2.3.4 Amorphophallus bulbifer (Roxb.) Blume..................................................................22
2.3.5 Amorphophallus coaetaneus S. Y. Liu & S. J. Wei................................................... 25
2.3.6 Amorphophallus corrugatus N. E. Br....................................................................... 28
2.3.7 Amorphophallus dunnii Tutch.................................................................................. 31
2.3.8 Amorphophallus kachinensis Engl. & Gehrm......................................................... 33
2.3.9 Amorphophallus kiusianus (Makino) Makino........................................................ 36
2.3.10 Amorphophallus konjac K. Koch.............................................................................. 38
2.3.11 Amorphophallus krausei Engl................................................................................... 40
2.3.12 Amorphophallus macrorhizus Craib.........................................................................43
2.3.13 Amorphophallus muelleri Bl. Blume (‘mulleri’)...................................................... 46
2.3.14 Amorphophallus paeoniifolius (Dennst.).................................................................. 50
2.3.15 Amorphophallus pygmaeus Hett..............................................................................54
2.3.16 Amorphophallus tenuistylis Hett.............................................................................. 56
2.3.17 Amorphophallus thaiensis (S. Y. Hu) Hett............................................................... 59
2.3.18 Amorphophallus tonkinensis Engler & Gehrmann................................................ 62
2.3.19 Amorphophallus variabilis Blume............................................................................64
2.3.20 Amorphophallus xiei H. Li & Z. L. Dao.................................................................. 67
2.3.21 Amorphophallus yuloensis H. Li.............................................................................. 70
2.3.22 Amorphophallus yunnanensis Engl.......................................................................... 72
2.4 Growth Cycles.......................................................................................................................77
2.5 Glucomannan Content in Various Amorphophallus Species............................................77
2.6 Hybrids...................................................................................................................................80
2.6.1 Hybridization in Amorphophallus..........................................................................80
Acknowledgements....................................................................................................................... 85
Appendix 2.I Successful Amorphophallus Hybridizations........................................................ 85
Appendix 2.II Failed Amorphophallus Hybridizations.............................................................. 87
References........................................................................................................................................ 94
5
2.1 Evolutionary and Genetic Aspects

The plant genus Amorphophallus belongs to the family Araceae, subfamily Aroideae, and
it is estimated that it has more than 200 species (Hetterscheid et al., 2012). This genus is
monocotyledonous and is distributed from tropical Africa throughout subtropical and
tropical Asia into the tropical western pacific and northeastern Australia (Hetterscheid
and Ittenbach, 1996). Among all the species of this genus about 70% have been found in
Southeast Asia (Boyce and Croat, 2011). The remarkable feature of this genus is its mor-
phology (see Figure 2.1) characterized by a solitary leaf rising from the tuber, consisting of
a vertical petiole and a horizontal leaf-blade (Hetterscheid and Ittenbach, 1996; Boyce et al.,
2012; for further details see Section 2.2).
There are several studies about this genus including its morphology, palynology and
biochemistry related to its odour. However, the morphology and palynology characters
and also the biochemistry of its volatile compounds are highly variable. The chromo-
some number of Amorphophallus is 2n = 24, 26, 28 or 3n = 39. Nearly 80% of the spe-
cies have 2n = 26, a small number exhibit 2n = 28 and some species are triploid with
3n = 39 (Wakabayashi, 1955). Some Amorphophallus with bulbils are triploid, with 39 chro-
mosomes (3n) that contribute to their suitability for vegetative propagation and vigor-
ous growth. The genome size of Amorphophallus is quite large, approximately 20 times
larger than the rice genome. C-value is the amount, in picograms, of DNA contained
within a haploid nucleus. A. johnsonii exhibits the largest genome size (1C = 15.83 pg),
A. bulbifer has a moderate genome size (1C = 9.28 pg) and A. prainii has the smallest
genome size (1C = 3.78 pg). In comparison, Oryza sativa (rice) has a genome size of only
FIGURE 2.1
Schematic representation of the morphology of Amorphophallus sp. (Adapted from Liu, P.Y., Konjac, China
Agriculture Press, Beijing, China, 2004; Chua, M., An investigation of the biology and chemistry of the Chinese
medicinal plant, Amorphophallus konjac, PhD thesis, University of Wolverhampton, Wolverhampton, UK, 2011.)
Botanical Background to Amorphophallus 7
0.50 pg/C (Chauhan and Brandham, 1985). Currently, a number of DNA techniques are
helping to determine the genetic relationships and evolution in this genus become more
clear through molecular phylogenetic analyses. A number of molecular markers have
been employed to determine relationships and to assess genetic variation in the genus.
The first study on the phylogeny of Amorphophallus – based on molecular markers by Grob
et al. (2002) – includes 46 Amorphophallus and Pseudodracontium species using chloroplast
matK and the trnL intron. The phylogeny showed that this genus can be divided into five
major clades according to their morphology. However, this study concluded that the rela-
tionship among phylogenetic clades remains unresolved and branches connecting these
clades are poorly supported. Later on, Grob et al. (2004) introduced a utility of FLint2
as a tool for phylogeny reconstruction in Amorphophallus. The FLint2 phylogeny was
shown to be largely congruent with chloroplast regions (rbcL, matK, and trnL). However,
there remained some limitations with unresolved issues of polytomies and lack of abil-
ity to provide enough non-conflicting informative characters to produce highly infor-
mative phylogeny. Subsequently, Sedayu et al. (2010) studied 69 Amorphophallus species
by combining gene-study data of trnL, rbcL in chloroplast and LEAFY (LFY) in nucleus.
The phylogeny showed major clades reflecting the biogeographical distribution and some
morphological synapomorphies. Five morphological characters showed relevant congru-
ence with the molecular phylogenetic results and lead to the following evolutionary state-
ments: (i) a non-sessile stigma may have evolved from a sessile one with several reversals;
(ii) pollen release through the rupturing of the connective evolved from pollen open-
ing by opening pores; (iii) unequally shaped main segments of the lamina evolved from
equally shaped segments three times in the Asian clades; (iv) simultaneous existence
of leaf (leaves) and inflorescence(s) evolved from alternating cycles of leaf and inflores-
cence; and (v) blue, purple, green, white and yellow fruits evolved from red/orange fruits.
However, this study on species-level relationships was based on a limited sample size
(30% of all species) with some under-represented clades and most of the relationships
remained with a high level of conflicting findings in the informative characters in the
tree. Claudel et al. (2017) enlarged the sampling to 157 species (70%), using nuclear (ITS1)
and plastid (rbcL and matK) regions. The combined phylogenetic trees (Figure 2.2) iden-
tified four main clades. The African clade (AFR-subgenus Afrophallus) contains all and
only African species and its unique seasonal cycle of both flowering, fruiting and leafing
in one and the same season. The Southeast Asian clade (SEA-subgenus Amorphophallus)
contains a majority of species from all over Southeast Asia (Indonesia, Philippines, east-
ern Malaysia), India, Indonesia to the Philippines and Australia (A. galbra). This group
is divided into (i) Paeoniifolius-Manta clade (12 species) that exhibits most of the red-
leafed seedlings and lack of offset development on the tuber; (ii) Pulchellus clade and (iii)
Pusillus clade with their monophyly being strongly supported by non-molecular charac-
ters such as their unique fruiting behaviour. The continental Asia I clade (CA-I-subgenus
Metandrium) includes species distributed in Asia with morphological support for part
of the clade by the sterile organs between the female and male zone. The continental
Asia II clade (CA-II-subgenus Scutandrium) includes mainly species from the Asian main-
land (India, southern China, Myanmar, Thailand and Indochina). This clade supports
the inclusion of the former genus Pseudodracontium in Amorphophallus (Hetterscheid and
Claudel, 2012).
In addition, genetic variation within the same species is interesting to study based
on different molecular markers; such as (a) microsatellite markers in A. paeoniifolius
(Santosa et al., 2007), A. konjac (Pan et al., 2012), A. bulbifer and A. konjac (Zheng et al.,
2013) and A. paeoniifolius (Santosa et al., 2017); (b) random amplified polymorphic
8
FIGURE 2.2
Combined phylogenetic tree of Amorphophallus with nuclear (ITS1) and plastid (rbcL and matK) region. (From Claudel, C. et al., Bot. J. Linn. Soc., 184, 32–45, 2017.)
Konjac Glucomannan
DNA (RAPD) markers in A. konjac (Wenbing et al., 2001), A. titanium (Poerba and
Yuzammi, 2008) and A. muelleri (Poerba and Martanti, 2008); and (c) amplified frag-
ment length polymorphism (AFLP) markers in A. paeoniifolius (Sugiyama et al., 2006)
and A. konjac (Pan et al., 2015).
Meanwhile, other markers were studied in relation to phylogenetic evolution, like
biochemical markers to identify the chemical composition of inflorescence odours of
Amorphophallus (80 species) by gas chromatography-mass spectrometry (GC-MS) (Kite
and Hetterscheid, 2017). When combined with the already described phylogenetic tree
(Claudel et al., 2017), the results showed that dimethyl oligosulphides were the dominant
odour compounds in Asian species. Fruity odours (1-phenylethanol derivatives) were the
dominant odour compounds in Metandrium clade, while anise odours (2-phenylethanol
derivative 4-methoxyphenethyl alcohol) are unique in Scutandrium clade. This study also
found that odour co-evolved with inflorescence colour, like rotting meat odours with
darker inflorescences. The evolution of some odour types is likely influenced by ecological
factors, the pressure of pollinator resources, etc.
Nevertheless, the present understanding of genetic relationships and evolution among
Amorphophallus species provides only baseline information. There are still some conflicts
in infrageneric classification and evolution based on complex morphological features and
additional factors such as plant distribution.
2.2 Taxonomy
The generic name Amorphophallus was proposed by Blume in 1825 (Bataafsche Courant,
nom. nud.) based on his encounter with A. paeoniifolius and later validated by him in
Decaisne (1834). The name is conserved against Thomsonia Wallich (1830) and Pythion
Martius (1831).
Schott (in Schott and Endlicher, 1832) was the first to present a classification of a num-
ber of Amorphophallus species, until then treated under Arum L., using the generic names
Candarum Reich. (nom. illeg.) and Pythonium Schott (nom. illeg.). Both names were pub-
lished superfluously.
Blume (1837) presented the first suprageneric and infrageneric classification of
Amorphophallus, treating it as part of the tribe Thomsonieae Bl. and dividing it into three
sections (Candarum (Reich.) Bl. (nom. illeg.), Adenophallus Bl. and Leiophallus Bl.).
Schott (1856) reduced Amorphophallus to the contents of Blume’s sect. Candarum.
Species of Blume’s other sections were transferred to Schott’s new genera Brachyspatha
Schott and Conophallus Schott. Schott also proposed the new genus Plesmonium. In sub-
sequent years (1857, 1858a, 1858b, 1860) Schott proposed several new genera, con-
taining species now accommodated in Amorphophallus, viz. Corynophallus, Hansalia,
Hydrosme, Rhaphiophallus and Synantherias. These genera were included in the tribe
Pythonieae Schott (1856, nom. illeg.) together with Pythonium, Anchomanes Schott and
Allopythion Schott and in 1860 Schott divided them over subtribes Amorphophallinae
Schott (Allopythion, Pythonium, Plesmonium, Rhaphiophallus, Synantherias, Brachyspatha,
Conophallus and Amorphophallus) and Hydrosminae Schott (Corynophallus, Hydrosme,
Hansalia and Anchomanes).
Hooker (1875) proposed the genus Proteinophallus to include A. rivieri Carr. (= A. konjac
K. Koch).
Engler (1876) reduced Conophallus, Proteinophallus and Brachyspatha to Amorphophallus

and Hansalia to Hydrosme. Subtribe Hydrosminae of Schott was entirely reduced to sub-
tribe Amorphophallinae except for Anchomanes being transferred to subtribe Pythoniinae,
now containing all genera with species possessing a sculptured appendix. Subtribe
Amorphophallinae is characterized by species with a smooth appendix. Both subtribes were
classified in tribe Amorphophalleae Engl. (nom. illeg., should be Thomsonieae Bl.). The lat-
ter tribe was classified with tribes Lasieae Engl. and Montrichardieae Engl. in subfamily
Lasioideae Engl.
In 1879, Engler produced a sectional classification of Amorphophallus. Three sec-
tions were recognized, viz. Candarum (Reich.) Engl. (nom. illeg., should be the autonym
Amorphophallus), Brachyspatha (Schott) Engl. (nom. illeg., mentioning sect. Leiophallus Bl. in
synonymy and by including A. muelleri the type of sect. Adenophallus Bl.) and Conophallus
(Schott) Engl. The remainder of his 1876 classification was not essentially changed. The sec-
tions were based on stylar length and shape of spathe and appendix.
In 1881, Engler described Hydrosme hildebrandtii, a species combining characters of
Corynophallus and Hydrosme. Engler therefore merged both genera, erroneously choos-
ing to use the name Hydrosme. Although this mistake was detected by Kuntze (1891)
and admitted by Engler (1893), he refused to change his decision and kept on using
Hydrosme illegitimately at the genus-level by including Corynophallus, the older name
of the two.
N.E. Brown proposed the genus Pseudodracontium in 1882 to accommodate a. o.
Amorphophallus lacourii Lind. & Andre.
Bentham and Hooker (1883) expanded Amorphophallus to include all genera mentioned
except for Thomsonia, Pseudodracontium, Synantherias and Rhaphiophallus. The same stance
was taken by N. E. Brown in 1901. Engler was long convinced that Hydrosme should be
kept separate from Amorphophallus on the basis of funicle morphology but finally gave
up in his last classification of the subfamily Lasioideae in Das Pflanzenreich (1911). In this
classification Amorphophallus, Thomsonia, Plesmonium, Pseudodracontium, Anchomanes and
Pseudohydrosme Engl. were accommodated in tribe Amorphophalleae. All other genera
were reduced to sections of Amorphophallus, to which were added several new ones. Of
these sections, Cundarum is an illegitimate name (see above) as well as (retroactively)
Conophallus since the type-species of Blume’s sections Leiophallus and Adenophallus are
included.
The last attempt to resurrect a genus from within Amorphophallus originates from
Ying (1991), publishing two new combinations of Taiwanese Amorphophallus species in
Hydrosme.
A few infrageneric names were added in previous and subsequent years viz. sect.
Dracontiopsis Engl. (1893, in Hydrosme, transferred to Amorphophallus by Engler in 1911), sect.
Podophallus Hook. f (1894), sect. Dysamorphophallus Engl. (1911), sect. Rapyogkos Engl. (1911),
sect. Cundaropsis Engl. (1911), sect. Interruptiflorus Engl. (1911), sect. Colliphallus Krause
(1924), subg. Metandrium Stapf (1924), sect. Titanum Nakai (1948) and sect. Exesispadix A.
D. Hawkes (1951). Engler’s final infrageneric classification remained unchanged, but for
the merger of sections Rhaphiophallus and Synantherias by Sivadasan (1989).
At the generic and suprageneric level, some major changes have been proposed. In 1985,
Bogner et al. proposed to sink Thomsonia in Amorphophallus (a case that was nomencla-
turally made possible by the accepted proposal of Nicolson et al. (1984) to conserve the
name Amorphophallus against Thomsonia). They also suggested sinking Plesmonium in
Amorphophallus. Hetterscheid (1994) finally presented arguments for the reduction of
Pseudodracontium as part of Amorphophallus and this step has been effected by Hetterscheid
and Claudel (2012).
At the level of tribe Thomsonieae, Bogner et al. (1985) proposed to remove Anchomanes
and Pseudohydrosme to another tribe (Nephthytideae Engl.) but leaving Thomsonieae in subf.
Lasioideae. Grayum (1984, 1990) was the first to treat the tribe Thomsonieae as part of sub-
fam. Aroideae but its position in it remained unclear. Bogner and Nicolson (1991) followed
Grayum but also refrained from further clarifying its position. Hetterscheid (1994) summa-
rized and added an extra argument for the tribe’s relation to subf. Aroideae and suggested
a sister-group relation to Arisaema Mart., based on preliminary cladistic analyses. More
recent analysis (molecular and morphological) indicates that Thomsoniae is a basal taxon
of a larger clade including genera of tribe Caladieae Schott (Cabrera et al., 2008; Cusimano
et al., 2011).
Molecular phylogenetic studies of Amorphophallus in its entirety (Grob et al., 2002, 2004;
Sedayu et al., 2010; Claudel et al., 2017) have all proven the monophyly of Amorphophallus in
the widest sense, including all satellite genera mentioned above by diverse authors, except
those transferred to other tribes than Thomsonieae. In Claudel et al. (2017) a first attempt is
presented at an infrageneric classification but only to the level of subgenera. The subgen-
era proposed are Amorphophallus (autonymic), Scutandrium Hett. & Claudel, Metandrium
Stapf and Afrophallus Hett. & Claudel. Further division of the genus at lower categorical
levels is hampered by low levels of statistical confidence at crucial nodes in the phylogeny.
A large body of literature exists introducing new species of Amorphophallus, as well
as floristic treatments (Thailand, China, Indochina, Africa, India, Java, Madagascar,
Taiwan) and partial taxonomic revisions (e.g., sect. Rhaphiophallus, sect. Conophallus, genus
Pseudodracontium). None of these three taxa were corroborated by the molecular results of
Claudel et al. (2017).
2.2.1 Amorphophallus: Genus Description

Amorphophallus species are geophytes, with the stem reduced to a storage organ usually
in the shape of a subterraneous tuber or more rarely a rhizome, either subterraneous (e.g.,
A. verticillatus Hett.) or just on the soil surface (e.g., A. hayi Hett.). The tuber may be globose,
subglobose or vertically elongated and then often branched, or as in one case only a chain
of tubers (A. coaetaneus S.J. Liu & S.J. Wei). Tubers may reproduce vegetatively by split-
ting up (e.g., A. titanum (Becc.) Becc. ex Arch.) or by producing few or many annual offset
tubers on their surface, more rarely by gradual offsetting. These offsets may be globose
(e.g., A. variabilis Blume), oval-elliptic (e.g., A. amygdaloides Hett. & Sizemore) or short or
long rhizomatous (e.g., A. albus P.Y. Liu & J.F. Chen). The size of the tuber varies from ca.
1 cm in diam. (e.g., A. ongsakulii Hett. & A. Galloway) to almost 1 meter (e.g., A. titanum).
Tubers of A. titanum may weigh up to 117 kg (pers. comm. Michael Neumann, from the
Botanical Garden Bonn, Germany)!
Annual growth of Amorphophallus species is cyclic (for more detail see Section 2.5).
The distinction between periods of the cycle are often distinct (e.g., a dormant period in
which there are no parts visible above ground) or less so (e.g., when leaves are longer
lived and only disappear when flowering starts). In tuberous species, the tuber is renewed
entirely during each full cycle and under optimal conditions will be bigger every time until
a species-dependent maximum is reached, after which the tuber usually splits.
Leaves develop directly on the tuber/rhizome and may be maximally ca. 4 cm high
in miniature species (e.g., A. ongsakulii) or reach a stunning 5 meter height in A. titanum.
In most species only one leaf develops per cycle. In a few clades of miniature species
(e.g., A. pulchellus Hett. & Schuit.) or species from the former genus Pseudodracontium (now
a clade in Amorphophallus, see Hetterscheid and Claudel, 2012; e.g., A. lacourii Linden &
Andre), more than one leaf may develop per cycle. Occasionally species develop more
leaves prior to a tuber split. This is often seen in A. paeoniifolius (Dennst.) Nicholson, for
example. And sometimes individual plants may develop more leaves when cultivated in
very rich soils or when considerable fertilizer is used (e.g., A. muelleri Blume).
The petiole (leaf stalk) may be smooth, or more rarely covered with velvety hairs (e.g.,
A. macrorhizus Craib), or variously set with smaller or larger warts (e.g., most forms of
A. paeoniifolius). The warts may be grouped around differently coloured spots and then
often imitate lichen (Claudel et al., 2019). The petiole may be of one colour (e.g., green,
purplish, brown) but more often it is variously ornamented with lines and/or spots of very
variable sizes and colour. Even within species, the colour pattern may vary considerably.
Quite often spots seem to imitate lichen and/or algae (Claudel et al., 2019) and may there-
fore imitate woodiness of the petiole. This may be a means to escape herbivory. In combi-
nation with the typical shape of the lamina (see below), the whole leaf thus often seems to
imitate young tropical sapling trees. The lower third part of the petiole often has a differ-
ent pattern of colours than the upper part.
The leaf blade is orientated perpendicular to the petiole. It is always dissected into
few or many smaller leaflets, superposed on a generally three-parted main division.
These three main divisions are in most species appr. of equal size and complexity but
in some groups, the anterior main segment may be less strongly developed than the
two lateral ones (e.g., A. lacourii), creating sometimes a pedate-like appearance, espe-
cially when the anterior segment carries one or few leaflets. Each main segment is
divided into higher order segments (the lamina is said to be ‘decompound’) and these
carry one, few or many leaflets. The diameter of the entire lamina may be as small as
ca. 10 cm (e.g., A. myosuroides Hett. & A. Galloway) or reach the enormous size of ca.
7 meters (A. titanum; pers. obs. Hetterscheid). The leaflets are shaped variously depend-
ing on the species (e.g., linear, lanceolate, narrowly to broadly elliptical or oval), their
surface usually being smooth but occasionally velvety hairy (e.g., A. atroviridis Hett.).
The colour of the upper surface is usually a shade of green. It may be a solid very-deep
velvety green (e.g., A. pendulus Bogner & Mayo), or occasionally variegated (e.g., spotted
as in A. lacourii, or striped along the midrib as in forms of A. pseudoharmandii Hett. &
Claudel), or feathered as in forms of A. taurostigma Ittenbach, Hett. & Bogner. Leaflets
of young sapling leaves of a few species may be purplish before turning green in older
plants (e.g., A. muelleri). In developing leaves all leaf segments and leaflets are pointed
upwards and the leaflets have their margins independently rolled inwards (involute
vernation). This character combination is the main genus-defining morphological trait
and underscores the monophyly of the genus. In several species groups, bulbils may
develop on branching points of the leaf lamina, which will produce new plants after
the leaf has died down (e.g., A. bulbifer (Roxb.) Blume, A. coaetaneus, A. muelleri, A. xiei
H. Li & Z.L. Dao, A. yuloensis H. Li).
The inflorescence of Amorphophallus species usually grows solitary from the tuber after
a period of dormancy except in a few species where it develops after leaves have devel-
oped (e.g., A. coaetaneus) or when the leaf is dying down (e.g., A. tuberculatus Hett. & V.D.
Nguyen). Occasionally more than one inflorescence develops alongside the leaves (e.g.,
A. pulchellus). Peduncle length varies between ca. 1 cm (e.g., A. terrestris) to 3–4 meters
in A. gigas Teijsm. & Binnend. and A. decus-silvae Backer & Alderw. In most cases the
peduncle is longer than the spathe, pushing the inflorescence well above soil surface
but some species develop a short peduncle and the inflorescence is sessile on the soil
surface, or almost so (e.g., A. paeoniifolius, A. yuloensis). Very few species show consider-
able variation in peduncle length (e.g., A. bulbifer). The surface and colour patterns of
the peduncle generally resemble those of the petiole but are often less prominent and/
or paler. In most species with sessile inflorescences, the peduncle may elongate strongly
after fertilization, pushing the (brightly coloured) fruits well in sight of potential dis-
tributers, notably birds. In one clade of small species, the long peduncle bows back to
the soil after fertilization and pushes the (dull coloured) fruits on the soil surface (e.g.,
A. pulchellus). In another clade of small species, the short peduncle does not (or hardly)
elongates at all after fertilization, leaving the (dull coloured) fruits close to soil surface
(e.g., A. terrestris Hett. & C. Claudel).
The flowering part of the inflorescence in Araceae is usually a pseudo-flower, consisting
of a variously shaped bract, called the spathe, situated at the base of an equally variable
axis with reduced flowers and often sterile organs, like (syn-)staminodes and pistillodes.
The axis is called the spadix. In Amorphophallus the variation in shape, colour and size of
spathe+spadix is bewildering. Size varies from very small, some 3–4 cm high in A. pusillus
Hett. & Serebryanyi, to the massive construct of A. titanum, which may reach up to 3 meters
in height!
The spathe in Amorphophallus is erect, at least the basal part of it. The basal part often
envelopes the lower part of the spadix and is usually closed with overlapping sides, form-
ing a vase-like or urn-like structure (e.g., A. muelleri). In two species the lateral sides are
not overlapping but fused, viz. A. elliottii Hook. f. and A. pusillus. In several species this
enveloping part is short or almost absent, exposing the base of the spathe (e.g., A. thaiensis
(S.Y. Hu) Hett.). The spathe base may be separated from the upper part (the limb) by a
constriction (e.g., A. kiusianus (Makino) Makino) or not (e.g., A. thaiensis). The outside of the
spathe base may be of one colour (often off-white, green or purple) or variously spotted.
The surface of the inside of the spathe base may be smooth but is more often verrucate,
with smaller or larger warts (e.g., A. paeoniifolius) or set with papillae, which are usually
hair-like, either short or long (e.g., A. angolensis (Welw. Ex Schott) N. E. Brown). The spathe
limb may be almost absent or more or less erect or horizontal and may be more or less flat
or convex. Its outer side may be of one colour (usually off-white, green or purple) or vari-
ously spotted and/or striped, the inside may be as the outside or paler and less complexly
patterned.
The spadix in Amorphophallus is differentiated in at least two different zones but most
often in three or even in four. The lower zone carries the female flowers, each flower
reduced to consist of only a pistil. Between the pistils sometimes hair-like staminodes
appear (e.g., A. cirrifer Stapf). Above the pistillate zone, usually a staminate zone follows
with strongly reduced male flowers, each flower consisting of 1–5 (or an indeterminate
number of) stamens. Occasionally hair-like staminodes are found between the functional
staminate flowers or within them (e.g., A. lanuginosus Hett.). Between the pistillate and
staminate zones, a sterile zone may exist, either devoid of any floral structure, or more
often set with staminodes (e.g., A. amygdaloides Hett. & Sizemore). Occasionally the neuter
flowers may turn out to be derived from pistils but this has not been unequivocally estab-
lished (e.g., A. longiconnectivus Bogner). Above the staminate zone, usually another sterile
part exists, called the appendix. The appendix consists of sterile stamen-derived tissue
and as such may be considered to be a single big synstaminodium. It may be smooth, or
variously set with staminodial (sub-) structures. In only one case, the appendix is truly
absent (A. margaritifer (Roxb.) Kunth). And in only one case the appendix is believed to be
secondarily entirely fertile (carrying functional stamens; e.g., A. coudercii (Bogner) Bogner).
The appendix is usually erect but may be bent horizontally (e.g., A. cirrifer Stapf) or may be
pendulous (e.g., A. pendulus Bogner & Mayo).
The pistils consist of an ovary, a style (but sometimes hardly differentiated) and a stigma.
The ovary is usually globose or depressed-globose and consists of 1–4 locules, each car-
rying 1 ovule. The style may be near-absent, short or very long, to at least 4 times the
height of the ovary. The stigma may be globose, depressed globose or disciform, lobed or
entire and very small to quite large, its diameter far outreaching the diameter of the style.
The stigma surface may be smooth or variously verrucate.
The fruits of Amorphophallus are usually fleshy, juicy berries, closely congested in a cylin-
dric infructescence. More rarely the infructescence is hemispheric or subglobose. The fruits
are globose or elliptic. The skin of the mature fruits is usually smooth and glossy but occa-
sionally verrucate and dull coloured (e.g., A. terrestris). The colour of most species is red to
orange but all species of one clade of the subgenus Metandrium have bright blue to purple
fruits (e.g., A. amygdaloides), whereas another group in this subgenus has mature fruits that
are yellow (e.g., A. atroviridis) or white (e.g., A. pygmaeus Hett.). Fruit size in Amorphophallus
varies from ca. 0.8 mm to 4–5 cm (in A. titanum). Fruits contain, dependent on the species,
1–4 seeds. Seed set usually requires cross-pollination but some species develop seeds apo-
mictically (e.g., A. bulbifer, A. kiusianus, A. muelleri and A. xiei.)
2.3 Glucomannan-Producing Species

The below presented descriptions are in several cases abbreviated and partly updated
from Li and Hetterscheid (2010) in Flora of China and Hetterscheid (2012) in Flora of Thailand.
2.3.1 Amorphophallus albus Liu & Wei

In: P. Y. Liu & J. F. Chen, J. S. W. Agric. Coll. 1984(1): 67.
Amorphophallus albus has a depressed globose tuber of up to 10 cm in diam., produc-
ing annually several, very long rhizomatous offsets. The leaf is solitary, up to ca. 1 meter
high. The petiole is green with greyish-green spots and whitish dots. The lamina is ca.
80 cm in diam. The leaflets are elongate-elliptic and up to ca. 12 cm long and 3 cm in
diam. The peduncle is up to ca. 80 cm long. The spathe is erect, up to 22 cm long, narrowly
ovate, not separated in base and limb, outside pale green, inside paler green. The spadix
is shorter than the spathe. The pistillate zone is 2–3 cm long, with closely packed pistils.
They have depressed globose, dark green ovaries, distinct thin whitish styles and stigmas
that are disciform and scabrate. A sterile zone is present above the pistillate zone, consist-
ing of large staminodes. The staminodes are irregular cushion-shaped or more rhombic
and flattened. Their surface is corrugated and off-white. The staminate zone is usually
longer than the pistillate, up to 5 cm. The stamens are dark orange-yellow. The appen-
dix is elongate fusiform-conical with a broadly rounded top. The surface is off-white and
usually irregularly verruculose (with small warts), the base often with a few staminodes,
similarly shaped as the ones in the neuter zone. The fruits are glossy orange-red and ovate
(Figures 2.3 through 2.7).
FIGURE 2.3
Amorphophallus albus: tuber with offsets. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.4
Amorphophallus albus: petiole detail. (Courtesy of C. Claudel.)
FIGURE 2.5
Amorphophallus albus: inflorescence. (Courtesy of C. Claudel.)
FIGURE 2.6
Amorphophallus albus: pistillate, sterile and staminate zones. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.7
Amorphophallus albus: fruiting. (Courtesy of C. Claudel.)
Distribution: China: Sichuan, Yunnan.

Ecology: in dry thickets of open forests; 800–1,000 m altitude.
NO T E :Amorphophallus albus belongs to subgenus Scutandrium. Although its morphology

is rather similar to that of A. krausei (including a staminodial zone and long, rhizoma-
tous offsets), molecular phylogenetic results (Gholave et al., 2016; Claudel et al., 2017) do
not resolve the relationships in this group.
2.3.2 Amorphophallus amygdaloides Hett. & Sizemore

In: Hetterscheid & van der Ham, Blumea 2001, 46(2): 253–282.
The tuber of this species is depressed globose and some 5–13 cm in diam. It produces
seasonally a few to many thick, fusiform offsets, which easily detach from the main tuber.
The single leaf has a petiole of up to 80 cm long and is ca. 2.5 cm in diam. Its surface is smooth,
mid to pale green with many circular or elliptic paler spots. The leaf blade is up to ca. 120 cm
across. The individual leaflets are elliptic-lanceolate, up to 18 by 6 cm, with an acuminate tip
and the upper surface is mid-green. The inflorescence appears solitary and never with a leaf.
The peduncle is up to 50 cm long and otherwise as the petiole. The spathe is elliptic, 21–24 cm
long and up to 20 cm in diam., strongly concave, hooded, top acute, the base only shortly con-
volute. The pouter surface of the spathe is pale green with whitish green spots, the inner sur-
face is uniformly whitish green but for a very small pinkish zone at the base. The inside of the
base is smooth. The spadix is shorter than the spathe, ca. 15 cm long, stipitate (meaning with a
short naked zone below the pistillate zone). The pistillate flower zone is cylindrical, 2 by 2 cm
with the pistils crowded. The ovary is depressed, mostly quadrangular in outline, 2 by 2.5 mm,
with a truncate top, whitish green, bilocular; the style is 1 mm long; the stigma is hemispheri-
cal, elliptic in outline, 2–3 mm in diam., shallowly or more distinctly bi- or trilobed. Above
the pistillate zone is a sterile zone with staminodes that is 1–1.5 cm long; the staminodes are
cushion-shaped, angular, creamy white. The staminate zone is cylindrical, 3–4 cm long, with
crowded staminate flowers. The stamens are creamy white. The appendix is fusiform-conical,
7–8 cm long and 2.7–3 cm in diam., sometimes laterally compressed, base constricted, apex
subacute, surface smooth, creamy white. Fruits glossy bright blue (Figures 2.8 through 2.13).
FIGURE 2.8
Amorphophallus amygdaloides: tuber with offsets. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.9
Amorphophallus amygdaloides: petiole detail. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.10
Amorphophallus amygdaloides: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.11
Amorphophallus amygdaloides: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.12
Amorphophallus amygdaloides: sterile and staminate zones. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.13
Amorphophallus amygdaloides: fruiting. (Courtesy of W.L.A. Hetterscheid.)
Distribution: Southwestern Thailand, Kanchanaburi province.

Ecology: not recorded.
NO T E : Amorphophallus amygdaloides is a member of a small group of species with broad

concave spathes and remarkably fruity scents (Kite and Hetterscheid, 1997, 2017), contrast-
ing well with the usual upsetting smells of a vast majority of species of Amorphophallus.
They (e.g., A. thaiensis, A. yuloensis, A. yunnanensis Engl.) all belong to subgenus Metandrium
and more particularly to a clear monophyletic group with the unique character of hav-
ing blue fruits, not found in any other group of Amorphophallus species. Amorphophallus
amygdaloides is the only species in this blue-fruiting group with a staminodial zone on the
spadix.
2.3.3 Amorphophallus asterostigmatus Bogner & Hett.

In: Bogner & Hetterscheid, Blumea 1992, 36: 467–475.
The tuber of A. asterostigmatus is depressed-globose or irregular and up to ca. 10 cm in
diam. and produces a few (or several short), elongate or subglobose offsets. The leaf is
solitary with a petiole of 60–70 cm long. The surface is smooth, silvery greyish-reddish
to greenish with scattered, suborbicular or elliptic, dark chocolate-brown spots. The leaf
blade is 50–100 cm across, with more or less elliptic leaflets that are 6–19 cm long and
2–5 cm wide, long, with the tip acuminate, adaxially dark green, abaxially paler. The inflo-
rescence develops without leaves and has a peduncle of 35–70 cm long, greenish to greyish
mauve with a few very dark brown spots. The spathe is erect, elongate ovate, as long as or
longer than spadix, base convolute, 15–21 by 7–13 cm, tip more or less cuspidate or apicu-
late. The outside of the spathe has a greenish margin, with a faint, pale purplish or reddish
brown tinge near the middle and the inside is cream or pale green, the base inside is green-
ish with or without irregular, pale purplish flushes and its surface is smooth. The spadix
is sessile or subsessile (lowermost 1.5–3 mm devoid of flowers), slightly shorter to slightly

longer than the spathe, 11–17.5 cm long. The pistillate zone is cylindric, 1–2.8 cm long with
the pistils congested. The ovaries are depressed-globose, 3.5–5 mm in diam., pale green,
unilocular; the style is conspicuously curved towards the spadix apex; the stigma is large,
deeply 2- to 4-lobed, irregularly star-shaped in upper view, margin strongly sinuous and
wavy. The staminate zone is cylindrical or slightly conical, 2.7–5 cm long with the staminate
flowers congested and entirely yellowish or with a reddish tinge at the apex. The appen-
dix is elongate-conical with a rounded apex, 6–11.5 cm long, largely smooth or with a
network of narrow, reticulated grooves, yellowish to cream-coloured, basal 1.5–2 cm with
conspicuous, irregularly elongate-rhombic, flat, rhombic to parallelogrammical, angulate
staminodes, often fusing upwards and resulting in the above described groove network.
Fruits unknown but most likely glossy red (Figures 2.14 through 2.18).
FIGURE 2.14
Amorphophallus asterostigmatus: tuber with offsets. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.15
Amorphophallus asterostigmatus: petiole detail. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.16
Amorphophallus asterostigmatus: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.17
Amorphophallus asterostigmatus: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.18
Amorphophallus asterostigmatus: pistillate and staminate zones. (Courtesy of W.L.A. Hetterscheid.)
Distribution: Central Thailand: Lop Buri prov., Saraburi prov.

Ecology: on slopes with limestone boulders.
N O T E : The inflorescence of A. asterostigmatus is most similar to A. longituberosus but

the latter has 2-, 3-, or 4-locular ovaries and a 2-, 3-, or 4-lobed stigma. Another con-
spicuous difference between these species is the shape of the tuber, which is depressed
globose in A. asterostigmatus but vertically elongate in A. longituberosus. Cushion-like,
rhombic staminodes at the base of the appendix are rare in Amorphophallus but may be
found in A. albus (see Section 2.3.1). Amorphophallus asterostigmatus belongs to subgenus
Scutandrium and has been placed close to A. schmidtiae Hett. & Galloway from Laos
(Claudel et al., 2017).
2.3.4 Amorphophallus bulbifer (Roxb.) Blume

In: Blume: Rumphia 1: 148. 1837. (Basionym: Arum bulbiferum (Roxb.), Fl. Ind. 3rd ed.: 510.
1832). Synonym: A. bognerianus Kumar & Jaleel, Aroideana 2009, 32: 136–141.
The tuber of A. bulbifer is subglobose to depressed-globose with distinctly swollen,
annulate rootscars. It may be up to 20 cm in diam. and never produces offsets. The leaf is
generally solitary, but more rarely a second leaf may develop during the season. The peti-
ole may be some 90 cm long and has a smooth surface with a dark green or brown-
ish green background colour with whitish, often rhombic, spots and sometimes linear,
white stripes. The base may be flushed pink. The leaflets are elliptic to elongate elliptic
to broadly lanceolate, up to 20 cm long, top acuminate, upper surface dull to glossy mid
green to dark green, often with a narrow pale pinkish margin (seedling leaves may be
dark green flushed with purple). On top of each of the main branching points of the
lamina a bulbil may develop, which is subglobose, brownish with small buds on the sur-
face. Bulbils may be up to 2 cm in diam. and produce new plants when buried in the soil.
The peduncle varies greatly in length and may be shorter (or less often much longer)
than the spathe, measuring from ca. 5 to ca. 80 cm long. It is otherwise similar to the
petiole. The spathe is erect and has quite a thick texture, measuring up to 40 cm with
the short basal part convolute but not clearly separated from the limb. The outside of the
spathe varies from green with many pale spots to entirely dirty pinkish and unspotted,
the inside of the base is usually dark pink, the limb is rarely pale green with some faint
spots but more usually uniformly pale to mid pink, the marginal zone usually paler than
the centre. The spadix may be slightly shorter to slightly longer than the spathe, varying
from ca. 10 to 35 cm long, and is sessile or stipitate. The pistillate zone is cylindric and
may be up to 5 cm long, with crowded pistils; the staminate zone is cylindric and some
2−3-times as long as the pistillate zone, measuring up to ca. 15 cm long. The appendix is
narrowly conical with a very thick wall and an obtuse apex, measuring up to ca. 15 cm
long. The surface of the appendix is minutely pitted and off-white, rarely dull brown.
Ovaries are depressed globose, up to 2 mm in diam., pale purplish or pale greenish, 1-
or 2-locular; style almost absent up to 1.5 mm long; stigma large, up to 3 mm in diam.,
hemispheric, very sticky, surface with a shallow central, often triradiate, depression, dirty
yellowish. Staminate flowers pale pinkish. Fruits broadly ovoid to elongate ovoid, bright
dark red (Figures 2.19 through 2.24).
FIGURE 2.19
Amorphophallus bulbifer: tuber. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.20
Amorphophallus bulbifer: leaf with bulbil. (Courtesy of C. Claudel.)
FIGURE 2.21
Amorphophallus bulbifer: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.22
Amorphophallus bulbifer: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.23
Amorphophallus bulbifer: pistillate and staminate zones. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.24
Amorphophallus bulbifer: maturing fruits. (Courtesy of W.L.A. Hetterscheid.)
Distribution: India (widespread, see Jaleel et al., 2012), Bangladesh, (Myanmar, acc.
to Jaleel et al., 2012 but no specimens mentioned).
NO T E :Amorphophallus bulbifer is a well-known species in circles of plant lovers. It has

been in cultivation in the western world for many decades (at least as early as the 1903)
because it tolerates quite low temperatures. A remarkable feature of this species is its trip-
loid chromosome number, 2n = 3x = 39 (Ramachandran, 1977; Chauhan and Brandham,
1985; Petersen, 1989; Anil et al., 2013). This effectively prohibits sexual reproduction in this
species. Its vegetative means of reproduction lie in the development of the foliar bulbils
and the nucellar development of clonal young plants from the pseudoseeds that develop
profusely without pollination (Brandham, 1983 [n.v.]). These characters are likewise found
in A. muelleri (see Section 2.3.13), the closest relative of A. bulbifer, and probably in A. xiei
too (but see Sections 2.3.4 and 2.3.20). Another result of the triploidy is disturbed pollen
development. Van der Ham et al. (1998) report a high percentage of diminished sterile pol-
len in A. bulbifer (some 75%) and likewise for A. muelleri, with the last one producing almost
no pollen at all. It is to be assumed that most, if not all, of the normal sized pollen grains
of both species are sterile.
2.3.5 Amorphophallus coaetaneus S. Y. Liu & S. J. Wei

In: Guihaia 6: 183. 1986. Synonyms: A. arnautovii Hett., A. pingbianensis H. Li &
C. L. Long.
Amorphophallus coaetaneus is a more or less evergreen species. The underground part
is a chain of depressed-globose, dark brown tubers, each one persisting after having
completed its cycle of leafing out and flowering/fruiting. A new tuber is then built up
on the older persistent one and so on, building a chain of tubers. Each tuber remains
in the chain a few seasons and then rots off. The chain may also branch. The leaf is
evergreen and persists some 3–4 years after development. The petiole is uniformly dark
green and may be up to 120 cm long. The lamina reaches ca. 100 cm across, with often a
bulbil developing on the main branching points. The leaflets are dark green, oblanceo-
late and ca. 20 cm long. The inflorescence develops alongside the leaf and its peduncle
reaches ca. 50 cm long. The spathe is erect and has a short convolute base, outside green
or dirty green flushed with dirty brownish purple or the upper part entirely brown-
ish purple; the inside is uniformly green or variously flushed with dirty dark purple.
The spathe is 7.5–15 cm long and the inside of the base is smooth or with a few scattered,
small, punctiform warts. The spadix is sessile, equaling the spathe or a little longer.
The pistillate zone is cylindric and 1–2 cm long, the pistils crowded. The staminate
zone is cylindric and 2.5–4.5 cm long, with stamens crowded. The appendix is narrowly
fusiform-conic, 5.5–14.5 cm long, sometimes slightly compressed, off-white to pale
green, smooth or shallowly corrugate, base sometimes with a few flattened staminodes,
separated by grooves, top acute. The ovary is green or pale whitish green, strongly
depressed, depressed globose, or subglobose, 1.5–4 mm in diam., 1- or 2-locular; style
straight or slightly upcurved; the stigma pale yellowish or pale greenish, flattened, dis-
ciform or nearly so, entire with a shallow, elongate, central depression or shallowly
2-lobed, or more distinctly bilabiate. The staminate flowers are off-white. The fruits are
dark blue (Figures 2.25 through 2.30).
FIGURE 2.25
Amorphophallus coaetaneus: tuber chain. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.26
Amorphophallus coaetaneus: leaf. (Courtesy of C. Claudel.)
FIGURE 2.27
Amorphophallus coaetaneus: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.28
Amorphophallus coaetaneus: pistillate and staminate zones. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.29
Amorphophallus coaetaneus: pistillate zone. (Courtesy of W.L.A. Hetterscheid)
FIGURE 2.30
Amorphophallus coaetaneus: fruiting. (Courtesy of R. McHatton.)
Distribution: South China: Guangxi (Guiping, Rongshui) & Yunnan; eastern and
north Vietnam.
Ecology: in moist forested valleys, along water, in thickets; 300–900 m altitude.
NOT E: A most unique species being the only one in the genus with a ‘rhizome’ of
chained tubers. The species is well-nested within the blue-fruited clade of subg.
Metandrium and the primitive-looking tuber chain may therefore well be an evolution-
ary novelty instead of a primitive character, which it looks like at first glance in the
light of the prevailing mode of tuber development in Amorphophallus (see Section 2.5).
The evergreen character of the species is shared with three other rhizomatous species
(A. verticillatus, A. hayi, A. rhizomatosus Hett., although all may be leafless when kept
dry [obs. in cultivation]).
2.3.6 Amorphophallus corrugatus N. E. Br.

Bull. Misc. Inform. Kew 1912: 269.
The tuber globose to subglobose, to 25 cm in diam., developing seasonal, long rhi-
zomatous offsets. The leaf appears solitary with the petiole reaching up to 95 cm, its
surface is shallowly, longitudinally ridged, the ground colour dirty white or pale green
with a very faint, pale brownish hue overlain with numerous tiny (and fewer large),
partly confluent, irregular, dark chocolate-brown or greyish spots. The lamina may
reach some 150 cm across, with the leaflets oblong or lanceolate, tip acuminate and up
to ca. 28 cm long., upper surface plain green. The inflorescence has a long peduncle of
up to 70 cm long, appearance as petiole. The spathe is ovate or elliptic-ovate, up to ca.
26 cm long, concave, top acute or obtuse, base shortly convolute. The spathe exterior is
pale greenish, greyish purple or white, sometimes to the base with whitish spots, or
with grey-green or pale olive-brown spots, with either only the margin purplish red or
also large parts reddish brownish. The interior of the spathe is pale greenish-whitish
with several irregular, purplish red spots and a purplish red margin, base within often
purplish red, or the utmost base whitish, surface smooth. The spadix is much shorter
than spathe, up to ca. 11 cm long, stipitate. The pistillate zone is cylindrical or slightly
obconical, to 3.5 cm long, pistils congested; the staminate flower zone is cylindrical to
fusiform-obconical, up to 4 cm long, flowers densely congested. The ovaries are globose,
depressed or subpyriform, 1–2 mm in diam., dark purple with pale whitish green base,
unilocular (occasionally bilocular); style slender, straight or more or less strongly curved
towards the spadix-axis, dark purple; stigma subapical or lateral, inconspicuous, usually
transversely bilabiate. Staminate flowers whitish, pinkish or pale purplish. The appen-
dix is slightly to distinctly stipitate, globose, ovate, elliptic or conical, obtuse or truncate,
1–5 cm long, surface with several irregular grooves (not associated with staminodes),
the longitudinal ones deepest, in between usually with a complex and dense pattern of
elongate, convoluted staminodes, separated by narrow grooves, the whole appearing
brain-like, yellowish white or pale greyish green, occasionally with a pale violet hue.
The fruits are orange (Figures 2.31 through 2.35).
FIGURE 2.31
Amorphophallus corrugatus: leaf. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.32
Amorphophallus corrugatus: petiole detail. (Courtesy of C. Claudel.)
FIGURE 2.33
Amorphophallus corrugatus: inflorescence. (Courtesy of C. Claudel.)
FIGURE 2.34
Amorphophallus corrugatus: spadix exposed. (Courtesy of C. Claudel.)
FIGURE 2.35
Amorphophallus corrugatus: pistillate and part of staminate zone. (Courtesy of C. Claudel.)
Distribution: Northern Thailand, N. Myanmar, SW China (SE Yunnan, Guangxi),

N. Vietnam. It is expected to occur also in Laos but no reports are known as yet
to the author.
Ecology: In primary evergreen forest on granite, in shade, 800–1,700 m altitude.
NO T E : Although this species bears great resemblance to A. yunnanensis and its allies, it
differs remarkably in developing orange-red fruits (instead of blue) and having long, rhi-
zomatous offsets (instead of short, subglobose ones). Also its phylogenetic position is very
different, being a member of subg. Scutandrium, whereas the yunnanensis-alliance is part
of subg. Metandrium (Claudel et al., 2017). A clear case of convergent evolution, possibly
forced by ecological circumstances because A. corrugatus is found in the same ecological

circumstances and geographical area as members of the yunnanensis-alliance.
The sister-species of A. corrugatus is A. kachinensis Engl. & Gehrm. (see Section 2.3.8).
The differences with this species however have turned out to be less significant than may
support species status. Basically the difference is the manner and intensity of the folding
of the appendix surface but several intermediate plants have been found, showing a mix of
defining characters of both species. It is therefore believed that both should be regarded as
one species, the priorable name would be A. kachinensis. This decision will await a separate
publication.
2.3.7 Amorphophallus dunnii Tutch

In: J. Bot. 49: 273. 1911. Syn. A. mellii Engler; A. odoratus Hetterscheid & H. Li.
The tuber is subglobose or depressed globose, to ca. 20 cm in diam., developing numer-
ous short, narrowly fusiform offsets annually. Leaf solitary with a green or greyish green
petiole with numerous elongate, confluent, pale green or brownish green spots, up to
60 cm long. The lamina is up to 100 cm in diam. with elliptic-lanceolate leaflets up to ca.
21 cm long, upper surface dark green. The inflorescence appears solitary and has a long
peduncle of up to ca. 60 cm long. The spathe is very broadly ovate and strongly concave
and up to some 25 cm long, with a very short basal convolute part. The outside is bright
pale green, basally with rounded white spots, these distally grading to whitish green; the
inside is similar but paler and the basal part is reddish purple. The surface of the inside
of the base is ridged-warty. The spadix is stipitate or rarely nearly sessile, slightly shorter
than spathe, up to ca. 20 cm. The pistillate part is cylindric, up to 2.5 cm long with the
pistils congested. The ovaries are ovary depressed and angulate, pale green, 2–3 mm in
diam., 2- or 3-loculed, with 1 basal ovule, the style is 0.5–1 cm long., the stigma yellowish,
flattened, ca. 2 mm in diam.; 2-, 3- or 4-lobed. The staminate zone is broadly fusiform and
up to 3 cm long. The appendix produces a scent of fresh carrots, is ivory-white, narrowly to
broadly conic, usually slightly dorsiventrally compressed, occasionally substipitate, vari-
able, hollow, up to 14 cm long, smooth or entirely echinate or with small warts, with occa-
sionally only the distal third part, base constricted and sometimes grooved. Fruits glossy
dark blue (Figures 2.36 through 2.40).
FIGURE 2.36
Amorphophallus dunnii: tuber with offsets. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.37
Amorphophallus dunnii: petiole detail. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.38
Amorphophallus dunnii: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.39
Amorphophallus dunnii: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.40
Amorphophallus dunnii: pistillate and part of staminate zone. (Courtesy of W.L.A. Hetterscheid.)
Distribution: China: Guangdong, Guangxi.
The blue berries and the ovate, concave spathe clearly group this species with
NO T E :
A. yunnanensis and allies in the blue-fruited clade of subg. Metandrium.
2.3.8 Amorphophallus kachinensis Engl. & Gehrm.

In: Pflanzenr. 48 (IV. 23C): 91. 1911. Syn. Amorphophallus bannaensis H. Li.
The tuber is depressed globose, up to ca. 30 cm in diam., developing offsets, probably
rhizomatous (n.v.). The leaf is solitary with the petiole up to ca. 100 cm long, dirty white
with dark green to reddish brown spots. The leaf blade is up to 100 cm across, with
elliptic leaflets of up to 15 cm long. Inflorescence solitary, peduncle up to 120 cm long,
surface as petiole. Spathe broadly elliptic-ovate, strongly concave, overarching the spa-
dix, up to ca. 30 cm long, basal part shortly convolute, outside green or greenish brown
with green spots or purplish red stripes and spots, inside dirty pale whitish-greenish or
with broad, longitudinal purplish stripes, or the reversed pattern, with a purplish back-
ground and several narrow, longitudinal whitish stripes, apex purple, base inside dirty
whitish green or pale purplish, with scattered shallow, tiny warts. Spadix, much shorter
than spathe, up to 18 cm long, stipitate. Pistillate zone slightly obconic or cylindric, up
to 5 cm long, pistils crowded or more loosely arranged. The ovaries are blackish purple,
globose or subpyriform, ca. 1.5 mm in diam., 1-loculed, the style is strongly curved and
dark purple; the stigma is very small, shallowly hemispheric or flattened, sometimes
more or less bilabiate. The staminate zone is obconic, occasionally fusiform, base often
constricted, to ca. 6 cm long, stamens congested and white or pale purple. The appen-
dix emits an unpleasant, rancid smell, is stipitate or occasionally sessile, dirty white or
with a narrow basal purplish margin above the stipe, ovoid or conic, occasionally nearly
globose, to ca. 7 cm long, sometimes with a few short longitudinal furrows at the base
but more often with a few deep, longitudinal grooves all over the length or with a few
shallow folds, or upwards more or less wrinkled, brain-like, glabrous or rarely shallowly
verrucate, base constricted, apex truncate, rounded, or rarely acute. Fruits orange red
FIGURE 2.41
Amorphophallus kachinensis: petiole detail. (Courtesy of C. Claudel.)
FIGURE 2.42
Amorphophallus kachinensis: inflorescence. (Courtesy of C. Claudel.)
FIGURE 2.43
Amorphophallus kachinensis: spadix exposed. (Courtesy of C. Claudel.)
FIGURE 2.44
Amorphophallus kachinensis: pistillate and part of staminate zone. (Courtesy of C. Claudel.)
Distribution: China: Yunnan (south and west); Laos; Myanmar, Kachin State; N. Thailand.
Ecology: in dense climax forests, on limestone rocks; 1,000–1,500 m altitude; flower-
ing March to May.
NO T E : See under A. corrugatus (Section 2.3.6).
2.3.9 Amorphophallus kiusianus (Makino) Makino

In: Bot. Mag. (Tokyo) 27: 244. 1913. Syn.: Amorphophallus konjac K. Koch var. kiusianus Makino;
A. hirtus N. E. Brown var. kiusianus (Makino) M. Hotta; A. sinensis Belval.
The tuber is depressed globose, up to ca. 20 cm in diam., with or without a few sessile, glo-
bose offsets. Leaf solitary, the petiole glossy, dirty olive-green or greyish green, with narrowly
oval or irregular whitish or very pale greenish spots and numerous tiny dark green dots, up to
ca. 65 cm long. The leaf blade is 60–90 cm across, with narrowly elliptic to lanceolate leaflets,
measuring up to 20 cm long of which the upper surface is bright green with a narrow pale violet
margin. The inflorescence is solitary with a long peduncle, coloured as petiole and 40–120 cm
long. The spathe is up to 25 cm long, outside dark greenish, greenish pinkish, or glossy dark
purplish brown, with small, whitish spots, margin with a narrow, reddish violet line, inside
pale pinkish with a purplish base, purplish and greenish, or entirely dark brown, with or with-
out a greenish margin, latter sometimes flushed pinkish, sometimes only medially pale green,
otherwise with rounded, whitish spots, base outside dark green or dark greenish brown, with
small, rounded whitish spots and blackish green veins. The spathe shape is triangular and
shallowly or clearly constricted between base and limb; limb at first oblique, then reflexing and
bending downward, margins reflexed or undulate; base inside dark purple, occasionally with
small, whitish spots and with numerous more or less distant, conic warts. The spadix is sessile
or subsessile, shorter than, equal to, or longer than the spathe and up to ca. 25 cm long. The pis-
tillate zone is slightly conical and up to 4 cm long with the pistils congested or slightly distant.
The ovaries are bright pale green, more or less obovoid, angulate in cross section, ca. 2 mm in
diam., 2-loculed, with the style very short. The stigma is pale greenish gray, shallowly or dis-
tinctly 2-lobed, sinuous, slightly oval in cross section, 1.5 mm in diam. Between the pistillate
and staminate zone is usually a short neuter zone, with purple hair-like staminodes of up to
1 cm long but sometimes this zone is almost missing but for a few staminodes. The staminate
zone is cylindric or slightly obconic, to 4.5 cm long with staminate flowers congested or distant
and pale yellow. The appendix is elongate conical, up to 20 cm long, blackish purple, smooth
or with a few hair-like staminodes at the base, these purple or whitish. Fruits at first green, then
turning carmine red or purplish pink and finally bright blue (Figures 2.45 through 2.49).
FIGURE 2.45
Amorphophallus kiusianus: leaf. (Courtesy of C. Claudel.)
FIGURE 2.46
Amorphophallus kiusianus: petiole detail. (Courtesy of C. Claudel.)
FIGURE 2.47
Amorphophallus kiusianus: inflorescence. (Courtesy of C. Claudel.)
FIGURE 2.48
Amorphophallus kiusianus: pistillate and part of staminate zone. (Courtesy of C. Claudel.)
FIGURE 2.49
Amorphophallus kiusianus: fruits maturing from lilac to blue. (Courtesy of C. Claudel.)
Distribution: China: Anhui, Fujian, Guangdong, Hunan, Jiangxi; Taiwan; southern

Japan: Kiushu.
Ecology: Shaded, semi-shaded, or sun-exposed places, plantations, secondary for-
ests, mixed bamboo and broad-leaved forests, orchards; 300–900 m altitude; flow-
ering from April to June, fruiting from May to July.
NO T E :The blue fruits indicate that A. kiusianus is part of subg. Metandrium. Its phyloge-
netic position within the blue-fruited clade remains unresolved (Claudel et al., 2017).
2.3.10 Amorphophallus konjac K. Koch

In: Wochenschr. Gärtnerei Pflanzenk. 1: 262. 1858. Synonyms: A. mairei H. Léveillé; A. nanus
H. Li & C. L. Long; A. rivieri Durieu ex Riviere.
Tuber brown, slightly glossy, depressed globose, up to ca. 40 cm in diam., seasonally pro-
ducing numerous long rhizomatous offsets with swollen apical part, which is often found
isolated after the connecting part to the main tuber has rotted away. The petiole may reach
150 cm long and has the background colour dirty whitish pinkish or dirty cream-coloured,
often nearly entirely covered by large, elongate, dark green confluent spots and smaller white
dots, or with numerous small, blackish green spots, very variable, with the surface glabrous
or with scattered punctiform warts at base. The lamina is up to 200 cm in diam. with ellip-
tical leaflets of up to 10 cm long and dull green on the upper surface. The inflorescence is
solitary and long pedunculate (rarely short). The peduncle may be up to 200 cm long and is
coloured as the petiole. The spathe measures 10–ca. 70 cm long, its base is coloured on the
outside dirty pale brownish with blackish green spots, or dirty pale whitish grayish with a
few scattered blackish green dots, near margin flushed with purple and on the inside maroon
with or without a paler whitish purplish zone above. The surface of the base inside is densely
warty and it is more or less separated from the limb by a shallow constriction. The limb is
erect, outside uniformly dark purplish brown, or with scattered blackish green spots, inside
uniformly dark brown, glossy, undulate and/or longitudinally folded, basal margin spread-
ing. The spadix is sessile, 15–110 cm long and longer than the spathe. The pistillate zone is
cylindric or narrowly conic, 2–11 cm long with the pistils congested or distant. The ovary is
depressed globose, whitish or pale pinkish, apex purplish, 2–4 mm in diam., 2- or 3-loculed.
The style is purplish, 1–5 mm long. The stigma is dirty yellowish brown; 2-, 3- or 4-lobed.
The staminate zone is cylindric, slightly fusiform, or slightly obconic, 2–12 cm long, staminate
flowers congested. The appendix narrowly fusiform-conic, 10–ca. 100 cm long, often laterally
compressed and with irregular, shallow longitudinal furrows, 10–85 × 1.5–6 cm, acute, dark
purplish brown or paler, densely rugulose, base often with several diamond-shaped, and flat-
tened staminodes. Fruits are orange-red (Figures 2.50 through 2.53).
FIGURE 2.50
Amorphophallus konjac: tuber with offsets. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.51
Amorphophallus konjac: leaf. (Courtesy of C. Claudel.)
FIGURE 2.52
Amorphophallus konjac: inflorescence. (Courtesy of C. Claudel.)
FIGURE 2.53
Amorphophallus konjac: pistillate and staminate zones. (Courtesy of W.L.A. Hetterscheid.)
Distribution: China, Yunnan. It is widely cultivated in China and Japan.

Ecology: in open situations or forest margins and thickets, secondary forests;
200–3,000 m altitude, flowering in April.
NOT E: It is unclear to date, which parts of the data above (morphology, distribution,
ecology) can be attributed to a ‘natural’ profile of the species, because it is unclear which
occurrences outside cultivated fields are assignable to naturally occurring individuals or
weedy escapees from cultivation. Amorphophallus konjac is one of the economically most
important species of the genus (the others being A. paeoniifolius and A. muelleri) and has
been domesticated many centuries ago. Philipp von Siebold brought plants from Japan,
which he named Arisaema konjac, the source of the present-day accepted species epithet
as established by Karl Koch. It is highly surprising to the author that to his knowledge no
known officially registered named cultivars are distinguished in this crop. Two names
are available for clones recognized in circles of ornamental plant lovers, viz. ‘Leo Song’,
a robust clone with very pale petiole and peduncle with only few and small darker spots
and ‘Nightstick’, a stunted clone with very dark petiole almost without spots and often
deformed inflorescences. In the light of the goal of this book, it may be suggested here
that it is high and about time that the Konnyaku-crop is described in terms of proper
cultivars, securing the possibility of choosing the most useful cultivars for particular
cultivation circumstances by farmers and for best choice of resources by glucomannan
producers.
2.3.11 Amorphophallus krausei Engl.

In: Engler, Pflanzenr. IV, 23C (Heft 48): 94. 1911. Synonyms: A. sutepensis Gagnep.; A. ximen-
gensis H. Li; A. pachystylis Hett.
The tuber is globose, sometimes slightly subcylindrical, with a deep central depression,
up to 25 cm in diam., seasonally producing several rhizomatous offsets. Leaf solitary, the
petiole up to ca. 200 cm long, pale green or mauve, at the base often pale pink or with a
reddish brown or reddish hue, with many, smaller and larger, elliptic, partly or almost
entirely confluent, to narrowly elliptic, blackish green or paler green or rarely reddish
brown spots and several small, white dots, the intensity of colours and the extension of
the pattern variable. The leaf blade is up to 160 cm across, with lanceolate leaflets of up to
48 cm length, plain mid green. The inflorescence appears solitary, with a long peduncle of
up to 100 cm and coloured as the petiole. The spathe is erect, boat-shaped, up to 40 cm long,
base short convolute. The spathe exterior is pale green, towards the base slightly darker;
the interior is pale yellowish green, with the base sometimes maroonish, and covered with
numerous small, slightly elongate or irregularly ridge-shaped, warts. The spadix is ses-
sile and nearly as long as spathe to distinctly shorter or slightly longer, up to 35 cm long.
The pistillate zone is cylindrical or slightly obconical, up to 5 cm long with pistils flowers
congested. The ovaries are globose or slightly depressed, 1.5–2 mm in diam., pale green,
occasionally pale magenta-purple near style base; style cylindrical, 1–2 mm long, green or
magenta-purple; stigma globose or subglobose, pale yellowish white, yellow or brownish,
ca. 1 mm, entire or with a shallow central depression or shallowly 2- or 3-lobed. Between
the pistillate and staminate zone is a sterile zone (rarely absent) of 0.5–2 cm long, densely
(or rarely distantly) set with staminodes which are ovate or diamond-shaped in cross-
section, rarely subglobose, ivory-white or creamy orange, occasionally flushed with pale
purple. Staminate zone up to 13 cm long, cylindrical-fusiform or slightly obconical, some-
times slightly laterally compressed, flowers congested. Staminate flowers ivory-white, the
lowermost flowers greatly enlarged and with reduced thecae, grading to the staminodes.
The appendix is short to elongate fusiform or fusiform-conical, up to 17 cm long, some-
times slightly laterally compressed, sometimes with a small stipe-like part, smooth, top
rounded or more or less acute, base occasionally stipe-like, sometimes with a few rounded
staminodes or staminodial remnants at the base, separated by small grooves, yellowish
white or pale green. Fruits are bright red (Figures 2.54 through 2.58).
FIGURE 2.54
Amorphophallus krausei: tuber with offsets. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.55
Amorphophallus krausei: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.56
Amorphophallus krausei: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.57
Amorphophallus krausei: pistillate, sterile and staminate zones. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.58
Amorphophallus krausei: fruiting. (Courtesy of C. Claudel.)
Distribution: Thailand: Chiang Mai, Chiang Rai, Nan, Kanchanaburi. China:

Yunnan; Myanmar (northern parts); Laos; Bangladesh.
Ecology: shaded to open, often fire-prone places in mixed primary evergreen/
deciduous forest and deciduous dipterocarp forest, often mixed with bamboo, on
granite bedrock, sometimes near streams. Lowlands to ca. 1,500 m altitude.
NO T E : The development of a sterile zone between pistillate and staminate zones with
gibbous staminodes has evolved several times independently in Amorphophallus (and in
Araceae as a whole) and as such is an unreliable character for suggesting evolutionary rela-
tionships. Of the species treated in this book A. krausei (subg. Scutandrium), A. albus (subg.
Scutandrium) and A. amygdaloides (subg. Metandrium) develop the aforementioned type of
sterile zone. The phylogenetic position of A. krausei relative to the other closest species in
subg. Scutandrium remains unresolved to date (Claudel et al., 2017).
2.3.12 Amorphophallus macrorhizus Craib

Kew Bull. 1912: 419. 1912. Synonyms: A. xiengraiensis Gagnep; A. longituberosus var. robustus
S. Y. Hu.
The tuber is vertically elongate, carrot-shaped, up to 35 cm long, unbranched or with few
branches in lower part. The leaf is solitary, with the petiole up to 110 cm long, its surface
densely, velvety hairy; the colour pattern is very variable, from almost entirely pale green
with a few elongate or long-linear, whitish spots, to reddish brown with many blackish
green, large confluent spots, sometimes almost entirely covering the petiole, many inter-
mediate patterns exist often suffused with reddish shades. The lamina is only moderately
to strongly divided, up to 110 cm across; the leaflets are elliptic to obovate, 10–36 cm long,
with the upper surface dark, velvety green with a narrow reddish purplish margin, the
lower surface greyish green, hairy like petiole. The inflorescence is solitary, long peduncu-
late. The peduncle is like the petiole and to 125 cm long. The spathe is erect, deeply boat-
shaped, up to 25 cm long, base convolute or nearly entirely open, grading into the limb or
slightly constricted at the apex, lower margins of the limb often strongly recurved, exterior
pale green with some small, darker green spots at the base, or with small, whitish spots
all over, the margin often suffused with a purplish hue, top dark green; the interior is pale
green with or without a purplish margin and sometimes with indistinct small whitish
spots all over and the basal part is green or dirty maroon, chequered with pale green or
entirely pale maroon, the latter colour sometimes extending over nearly the entire spathe.
The inside of the basal parts has the veins impressed, groove-like, in between with very
tiny, punctiform warts. The spadix is sessile or very shortly stipitate, slightly to distinctly
longer than spathe and up to 30 cm long. The pistillate zone cylindrical or slightly conical,
up to 5 cm long, pistils congested, rarely a small naked zone at the apex or a small zone
with flask-shaped pistillodes. The ovaries are depressed, prismatic or diamond-shaped, up
to 3 mm in diam., entirely bright pale green or with a blackish purple apex; style absent or
very short; the stigma is 2-, 3- or 4-lobed, oval in outline, 1 by 1–1.75 mm, greyish or pale
yellowish brown. The staminate zone is fusiform-conical, up to 12 cm long, staminate
flowers congested. The appendix is elongate fusiform, subulate, sometimes very feeble,
5–21 cm long, often compressed and slightly sinuous, the base often with several, scat-
tered, flexuous, ca. 1–3 mm long, thin, purple-brown hairs, otherwise smooth, dirty (yel-
lowish) green, blackish brown or lead-grey, the base often white and sometimes more or
less stipe-like. Fruits are bright red (Figures 2.59 through 2.63).
FIGURE 2.59
Amorphophallus macrorhizus: tuber. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.60
Amorphophallus macrorhizus: leaf. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.61
Amorphophallus macrorhizus: petiole detail. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.62
Amorphophallus macrorhizus: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.63
Amorphophallus macrorhizus: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
Distribution: Thailand: Mae Hong Son, Chiang Mai, Chiang Rai, Lampang, Tak,
Udon Thani.
Ecology: dry deciduous dipterocarp-oak forests, on shale or granite in very hard soil;
300–1500 m altitude.
NOTE: Amorphophallus macrorhizus is a member of subg. Metandrium and teams

up with several species all bearing carrot-shaped tubers and some of which also
develop hair-like staminodes on the spadix, albeit in much higher numbers than
A. m acrorhizus. The development of elongate, carrot-like tubers is not an evolutionary
unique feature in Amorphophallus and developed at least once in three of the subgen-
era (Claudel et al., 2017).
2.3.13 Amorphophallus muelleri Bl. Blume (‘mulleri’)

In: Blume: Rumphia, 1: 143. 1837. Synonyms: A. planus Teijsm. & Binnend., A. oncophyllus
Prain, A. burmanicus Hook. f., A. carnosus Engl., A. timorensis Alderw., A. erubescens
Hett.
The tuber is globose or depressed-globose, up to 30 (or more?) cm in diam., without
annual offsets. The leaf is solitary, occasionally two on one tuber. The petiole is smooth,
up to 180 cm long, green, olive-green, brownish green or almost black, with numer-
ous, large, elongate-elliptic, diamond-shaped or stripe-like, pale green spots and, some-
times, with an additional high number of small, pale green, rounded dots. The lamina
is highly dissected, up to 200 cm across, with bulbils developing on the major points of
branching and these are depressed, rounded or elongate, 0.5–6 cm diam. The leaflets
are lanceolate or elliptic lanceolate, up to 40 cm long, green or dark green with a nar-
row, whitish or pinkish red margin, especially when young. The inflorescence is soli-
tary, long or (more rarely) short pedunculate, with the peduncle being 10–60 cm long.
The spathe is broader than long, rarely slightly the reverse, very broadly triangular or
transversely elliptic, up to ca. 35 cm long, leathery, base strongly convolute and slightly
or clearly separated from the limb by a constriction. The limb is erect upon opening and
then spreads to almost horizontally, with the margin variously reflexed. The outside of
base of the spathe is pale green or pale dirty pinkish usually with transversely elongate,
whitish spots and few small, blackish green dots, whereas the limb usually has large,
more or less isodiametric white spots. The inside of the base of the spathe is dark pink,
bright pale pink or pale yellowish pink, upwards grading to purplish, pink, dark pink,
dark brownish pink or brownish with dirty pale greenish and dirty pale brown, trans-
versely oval whitish spots, or more rarely the entire interior of the spathe reddish pink.
The inner surface of the spathe base is nearly smooth or with numerous small, elongate
warts, these often confluent. The spadix is sessile or stipitate, equal to or longer than
spathe, up to 40 cm long. The pistillate zone is cylindrical or slightly fusiform, 1.5–10 cm
long, with pistils congested or more loosely arranged. The ovaries are short stipitate,
depressed, 3–3.5 mm in diam., the lower half reddish purple, upper half pale pink, 2- or
3-locular; the style is short, 0.3–1 mm long. Stigma large but diameter always slightly
smaller than ovary, circular, quadrangular or slightly oval in cross-section, flattened,
subhemispherical or hemispherical, 1–2 mm across, with a shallow, elongate or three-
rayed central depression or shallowly 3- or 4-lobate, margins reflexed, occasionally
with two or three, dirty pale yellow. The staminate zone is cylindrical, obconical or
fusiform, up to 9 cm long, in large specimens usually laterally compressed, flowers
congested. The appendix is narrowly or very broadly fusiform-conical, 3–22 cm long,
laterally compressed to various degrees, surface pitted with numerous small, puncti-
form depressions and with or without some irregular, larger shallow depressions, base
with staminodes intermediate between stamens and the appendix-wall, pale pinkish,
yellowish or pale brownish. Fruits are red (Figures 2.64 through 2.71).
FIGURE 2.64
Amorphophallus muelleri: old leaf with bulbils. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.65
Amorphophallus muelleri: petiole detail. (Courtesy of C. Claudel.)
FIGURE 2.66
Amorphophallus muelleri: flowering plant from Thailand, Kanchanaburi. (Courtesy of Yamazaki.)
FIGURE 2.67
Amorphophallus muelleri: inflorescence. (Courtesy of C. Claudel.)
FIGURE 2.68
Amorphophallus muelleri: spathe view of inner surface. (Courtesy of C. Claudel.)
FIGURE 2.69
Amorphophallus muelleri: spadix exposed. (Courtesy of C. Claudel.)
FIGURE 2.70
Amorphophallus muelleri: pistillate zone. (Courtesy of C. Claudel.)
FIGURE 2.71
Amorphophallus muelleri: fruiting. (Courtesy of C. Claudel.)
Distribution: Thailand: Chiang Mai, Tak, Kanchanaburi; India: Andaman & Nicobar
Islands; Myanmar; Indonesia: Sumatra, Java, southern Borneo, Flores, Timor.
Ecology: ruderal and open secondary seasonal forests.
NO T E : The correct application of the name A. muelleri has been chaotic since Engler (1879)
created a mixture of true A. muelleri and probably a west Javan species (A. annulifer Hett.)
and perpetuated this in his Lasioideae treatment in 1911. Several authors duplicated this
in flora’s, etc., so when Prain discovered a plant of A. muelleri on Great Coco Island, he
considered it a different species than the one from Java and named it A. oncophyllus (based
on the bulbils on the leaves). Van Hasselt’s drawing of the original plant (used by Blume
to create A. muelleri), leaves no doubt about it being the same species as meant by Prain.
Jaleel et al. (2012) decided to ignore this and accepted that the name A. oncophyllus repre-
sents a different species and even restricted its occurrence to the Andaman Islands, there-
fore claiming it to be an Indian endemic. This is impossible in the light of the fact that
Prain’s type specimen of the name of this species is from Great Coco Island, belonging to
Myanmar. Specimens of A. muelleri from eastern Thailand often enough show characters
that are similar to plants from the Andamans and the (taxonomically irrelevant) difference
that remains basically is the darker reddish-brownish colour of the spathe of the latter
specimens. Jaleel et al. (2012) also uphold the species A. carnosus, based on a specimen that
only differs from the other muelleri on the Andamans by having a somewhat aberrantly
long attenuated appendix. This is also found, be it lesser pronounced, in specimens from
Thailand. Also the type of A. carnosus does not show this character, nor is it mentioned in
the protologue, so their plant is just a singly occurring, aberrated A. muelleri.
2.3.14 Amorphophallus paeoniifolius (Dennst.)

In: Nicolson: Taxon 26 (2/3): 338. 1977. Basionym: Dracontium paeoniifolium Dennstedt, Schl.
Hort. Ind. Mal. (1818) 13/21/38 (‘paeoniaefolium’). Synonyms: A. campanulatus Decne (non
Roxb.); A. paeoniifolius var. campanulatus (Decne) Sivad.; A. sativus Bl.; A. decurrens (Blanco)
Kunth; A. rex Prain; A. malaccensis Ridley; A. campanulatus var. australasicus (‘australasica’)
Maiden; A. dixenii K. Larsen & S. S. Larsen.
Tuber depressed globose, up to ca. 50 cm in diam. Rootscars prominently thickened,
annulate, offsets produced every season, these thick rhizomatous, up to ca. 10 cm long.
Leaf solitary or paired, rarely more; petiole up to ca. 2 m long, background colour pale
to dark green or blackish green, usually with large and small pale blotches and numer-
ous tiny dark dots, the large blotches often confluent, especially near the base, surface
near smooth, shallowly corrugate to strongly echinate-verrucate. The lamina is highly
dissected, up to ca. 3 m. in diam. The leaflets are orbicular, oval, ovate, obovate, elliptic,
elongate elliptic, elliptic-oblong, acuminate and up to 35 cm long with the upper surface
mid green. The inflorescence is short pedunculate and sits more or less on the soil surface.
The peduncle during flowering is 3–20 cm long, usually paler and smoother than petiole
but elongates enormously during fruit set becoming as long 100–200 cm, the surface turn-
ing uniformly pale brown. The spathe is campanulate, broader than long, up to ca. 45 cm
long, base and limb often separated by a shallow constriction, limb spreading, strongly
undulate, base outside vary variable, background colour ranging from pale green to dark
brown, usually with large and small, circular paler spots, base inside lower part deep
maroon, upper zone dirty whitish or very pale pinkish, limb outside as base but with
more prominent maroon flushes, especially near the margin, limb inside usually glossy
dark maroon, base within densely warty with the warts variable, mostly conical, fleshy.
The spadix is sessile, shorter or longer than spathe, up to ca. 70 cm long. The pistillate zone
is cylindric, 3–25 cm long with the pistils congested or slightly distant. The staminate zone
is cylindric or strongly obconic and roofed against the expanded base of the appendix, up
to 15 cm long with the staminate flowers congested. The appendix very variable, inflated,
1.5–ca. 30 cm long, 1.2–30 cm in diam. (slightly above the base), globose, depressed glo-
bose, oval or triangular conic (pyramidal), smooth or with various folds and/or irregular
shallow depressions, base often with flattened, staminodial structures, top obtuse or acut-
ish, surface minutely granulate, glossy dark maroon, rarely pinkish or yellow, giving off
a stench of rotting meat. The ovaries are depressed, circular in cross-section, 3–5 mm in
diam., 2- or 3-locular, entirely pale green or largely maroon with a whitish base. The style
is long and slender, up to 20 mm long, maroon. The stigma is large, oval or triangular in
cross-section, 4–7 mm in diam., often strongly laterally compressed, shallowly or deeply
2- or 3-lobed, pale or deep yellow. The staminate flowers are off-white. Fruits are bright red
FIGURE 2.72
Amorphophallus paeoniifolius: tuber. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.73
Amorphophallus paeoniifolius: leaf. (Courtesy of unknown.)
FIGURE 2.74
Amorphophallus paeoniifolius: petioles appearing from same tuber. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.75
Amorphophallus paeoniifolius: petiole detail. (Courtesy of C. Claudel.)
FIGURE 2.76
Amorphophallus paeoniifolius: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.77
Amorphophallus paeoniifolius: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.78
Amorphophallus paeoniifolius: pistillate zone. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.79
Amorphophallus paeoniifolius: fruiting. (Courtesy of unknown.)
Distribution: Madagascar, eastwards via India to Malesia (biogeographical region),

south China, Indochina, Polynesia, northern Australia.
Ecology: in almost all imaginable secondary conditions, either secondary forest or
highly disturbed areas, in dappled shade or fully exposed grounds, 0–700 m alti-
tude; flowering April-June (SW India); March-June (Thailand); May (NE India);
May (West Malaysia); June (West Malaysia); June (Philippines); July (N Vietnam);
Oct. (Fiji); November (Java).
NO T E : Amorphophallus paeoniifolius has been widely cultivated throughout Asia. The spe-
cies seems to become weedy quite easily after escaping from cultivation. It is therefore
certain that parts of the descriptive, geographical and ecological data relate to characters
and data that deal with weedy escapes from cultivation and are not part of the natural
‘evolutionary’ make-up of this species. Suresh et al. (1983) have attempted to separate the
wild (as A. paeoniifolius var. paeoniifolius) and cultivated (as A. paeoniifolius var. campanula-
tus) aspect taxonomically and thus used the concept of ‘taxon’ as used for wild plants for
domesticated plants and consequentially using infraspecific Linnean epithets to express
their ideas. It is however clear that, at least in the Malesian area, there is no straightforward
distinction in morphology between cultivated and wild aspects of this species, provided it
is at all possible to make the distinction in the first place without thorough ethnobotanical
studies. Local taxonomic solutions may be looked for to distinguish the cultivated aspect
in terms of cultivars, for which no existing taxon names are necessary and warranted
other than folk names or accurately established cultivar epithets. A useful study of wild
and weedy aspects of this species should be based on concepts in use for domesticated
plants and described in the nomenclatural code for cultivated plants (Brickell et al., 2016)
and not the nomenclature code for algae, fungi and plants.
2.3.15 Amorphophallus pygmaeus Hett.

In: Blumea 39(1–2): 271. 1994.
A very small species with a short elongate tuber, or more subglobose, often somewhat
irregular, rarely (young tubers) depressed globose, rarely branched, ca. 2–6 cm long. Leaf
solitary with the petiole 10–40 cm long, smooth, uniformly reddish brown or pale olive-
brown. The leaf lamina moderately dissected, to 55 cm across, with the leaflets elliptic
lanceolate, obovate or oblong, to 13 cm long, upper surface very dark velvety bluish green
with a reddish margin, lower surface flushed iridescent purple-red. The inflorescence
appears solitary, long pedunculate. The peduncle is coloured like the petiole or paler, to
42 cm long. The spathe is erect, base convolute and with rather strong concave sides, to
10 cm long, outside and inside creamy white or pinkish white, base inside dirty brown-
ish red and with scattered or numerous fleshy, shortly elongate, irregularly branched or
laterally flattened warts. The spadix is sessile, longer than the spathe, up to 14.5 cm long.
The pistillate zone shortly cylindrical, up to 1.5 cm long, pistils congested or slightly
distant. The ovaries are depressed, diamond-shaped, angular or circular in cross-sec-
tion, up to 1.5 mm in diam., pale green, unilocular; style short; the stigma hemispherical
or depressed, very shallowly multilobed and with a shallow central depression, dirty
creamy white. The staminate zone is fusiform, fusiform-obconical or slightly conical, up
to 5 cm long, staminate flowers congested, white with a pale greenish hue. The appendix
is elongate-fusiform, smooth, very pale green or creamy white, top obtuse, up to 8 cm
long. Fruits white (Figures 2.80 through 2.84).
FIGURE 2.80
Amorphophallus pygmaeus: tuber. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.81
Amorphophallus pygmaeus: leaf. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.82
Amorphophallus pygmaeus: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.83
Amorphophallus pygmaeus: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.84
Amorphophallus pygmaeus: pistillate and staminate zones. (Courtesy of W.L.A. Hetterscheid.)
Distribution: Thailand: Prachuap Khiri Khan, Saraburi.

Ecology: medium-shaded, humus-filled crevices and pockets on limestone; 50–500 m
altitude.
NO T E :Amorphophallus pygmaeus belongs to a small, well supported clade in subg.

Metandrium, of which all species are small, possess elongate tubers, deep emerald green
leaves with pinkish margins, white spathes and white fruits.
2.3.16 Amorphophallus tenuistylis Hett.

In: Blumea 39(1–2): 279. 1994.
The tuber is elongate, carrot-shaped, up to 16 cm long, unbranched. The leaf is sol-
itary with the petiole up to 80 cm long, smooth, largely pale green with narrowly
elongate-elliptic, pale to dark green spots, blackish green at the base, or largely dirty
pale grey with elongate-elliptic, dark olive-green spots, often confluent, and some sim-
ilar small dots near the base, or almost entirely dark olive-green with numerous small,
greyish punctate spots. The leaf lamina is up to 100 cm across, with the leaflets elliptic,
elongate-elliptic or lanceolate, up to 12 cm long. The inflorescence is solitary with the
peduncle up to 100 cm long, greyish to pale grey with elongate, often confluent, olive-
green or dark brown spots and scattered small whitish dots. The spathe is erect, elliptic-
lanceolate to elongate-triangular, up to 40 cm long, base convolute and separated from
the limb by a shallow constriction, apex acute, margin strongly sinuous, sometimes
spirally twisting along the midrib, exterior appearance as petiole but limb flushed
maroon, interior base pale to dark maroon, limb dark maroon, or dirty green with
purple-brown flushes, darker near the margin, or dirty pale grey with orbicular, dark
green spots around the midrib and the remainder dark purple, base within densely
clothed with short and long, more or less fleshy, elongate, hair-like warts, sometimes
with a few irregular branches. The spadix is sessile or stipitate, shorter to distinctly
longer than spathe, 7–50 cm long. The pistillate zone cylindrical, flowers slightly or dis-
tinctly distant, up to 6.5 cm long, with or without a very short naked zone at the apex.
The ovaries are depressed, oval in outline, up to 4 mm in diam., more or less bright
pale green; style slender, dark brown or pale purple; stigma obconical, 1.5–2 mm in
diam., entire or shallowly, or distinctly 2- or 3-lobed, off-white or pale yellow. A short
sterile zone between pistillate and staminate zone may be present and then with dark
purple brown, angulate, flattened staminodes. The staminate zone is cylindrical or
slightly conical, up to 11 cm long, flowers congested. The staminate flowers are off-
white or purplish and the ones at the base of the zone are often enlarged. The appendix
is narrowly elongate, acute, in smaller inflorescences even mouse-t ail-like, up to 32 cm
long., lower part sometimes warty, otherwise smooth, brownish grey. Fruits are orange
red (Figures 2.85 through 2.89).
FIGURE 2.85
Amorphophallus tenuistylis: tuber. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.86
Amorphophallus tenuistylis: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.87
Amorphophallus tenuistylis: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.88
Amorphophallus tenuistylis: pistillate zone. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.89
Amorphophallus tenuistylis: fruiting. (Courtesy of W.L.A. Hetterscheid.)
Ecology: Open mixed bamboo & deciduous forest, on limestone; ca. 200 m altitude.
NO T E :Amorphophallus tenuistylis belongs to a small clade of species in subg. Scutandrium,

with long tubers, A. longituberosus (Engl.) Engl. & Gehrm. and A. albispathus Hett. Characters
in common are the long warts in the spathe base and the three-locular ovary and the
enlarged lower staminate flowers. A sister subclade contains species that look generally
much more like A. tenuistylis and it may well turn out to be closer to those species than
indicated by the present phylogenetic analysis, with moderate confidence levels for the
grouping of the three above-mentioned species.
2.3.17 Amorphophallus thaiensis (S. Y. Hu) Hett.

In: Flora of Thailand 11(2): 178. 2012. Basionym: Thomsonia thaiensis S.Y. Hu
The tuber is subglobose, up to 12 cm in diam., with numerous large, elongate-f usiform
offsets. Leaf solitary; petiole up to 60 cm long, smooth, olive-brown with indistinct
slightly paler shades and scattered, dirty pale grey, elongate-elliptic or somewhat
irregular spots. The leaf lamina is up to ca. 60 cm across with relatively few leaflets.
The leaflets are elongate-elliptic, short acuminate, up to 18 cm long, upper surface
dark green, somewhat glossy. Inflorescence solitary, long-pedunculate; peduncle up to
40 cm long, smooth, reddish brown with scattered, circular or rhombic, dirty whitish
or dirty pale reddish brown spots. The spathe is broad, concave, boat-shaped, up to
22 cm long, base shortly convolute, top acute, outside dirty greenish with or without
a strong brownish purplish flush at the base or more extensively, especially in imma-
ture spathes, upper part sometimes dark green, with several scattered or congested,
distinct or indistinct, small or large, partly confluent, dirty whitish or pale green spots
in the lower part or all over, inside pale green, with a faint purplish hue near the base,
lower margins narrowly purplish. The spadix is stipitate, shorter than the spathe, up
to 10 cm long. The pistillate zone is cylindrical, up to 1.5 cm long, flowers congested.
The ovaries are obliquely depressed, 2–4 mm in diam., bright pale green or maroon,
unilocular; the style is 2–3 mm long, straight or curved towards the spadix, bright pale
green or maroon; the stigma is strongly depressed, bilabiate with an elongate central
depression, dirty pale yellowish. The staminate zone is cylindrical, up to 2.5 cm long,
top slightly laterally compressed, flowers congested; staminate white. The appendix is
ovoid to shortly conical, 2–6.5 cm long, not or slightly laterally compressed, with or
without one or two broad, longitudinally elongate depressions, top broadly obtuse, sur-
face smooth, dirty white, with or without a pale purple flush. Fruits are glossy bright
blue (Figures 2.90 through 2.94).
FIGURE 2.90
Amorphophallus thaiensis: tuber with offsets. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.91
Amorphophallus thaiensis: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.92
Amorphophallus thaiensis: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.93
Amorphophallus thaiensis: pistillate zone. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.94
Amorphophallus thaiensis: fruiting. (Courtesy of W.L.A. Hetterscheid.)
Distribution: Thailand: Mae Hong Son, Chiang Mai.

Ecology: In humus-filled limestone pockets, in soil on slopes, in medium shade.
NO T E :With its blue fruits, A. thaiensis is yet another member of the blue-fruited clade of
subg. Metandrium. It mostly resembles A. yunnanensis but differs in the much smaller, later-
ally compressed, bilabiate stigma and the much higher number of annual offsets.
2.3.18 Amorphophallus tonkinensis Engler & Gehrmann

In: Pflanzenr. 48 (IV. 23C): 87. 1911.
The tuber is depressed globose, up to 35 cm in diam. (or more?), rootscars slightly raised,
no offsets observed. The leaf is solitary, with the petiole up to 200 cm long, smooth, uni-
formly pale green or background pale greyish green with a clear bluish flush near base and
on subterranean part, and with whitish spots, these often broader than long, irregularly
shaped, and scattered over entire surface with short, longitudinal, blackish dots, these
in proximal part often raised. The leaf lamina is highly dissected, up to 200 cm across,
sometimes some nodes swollen and forming intercalary bulbils. The leaflets are up to
24 cm long, elliptic-lanceolate to lanceolate, long acuminate. The inflorescence is solitary,
long pedunculate, with the peduncle up to 50 cm long (or more?), coloured as petiole.
The spathe is up to 20 cm long (the sizes of the flowering parts are based on cultivated
specimens only and are likely larger in the field), with a shortly convolute base, strongly
concave, erect or arching over spadix apically, the outside of the base is green or brown
with a few transverse, whitish spots, its inside is whitish green or pale brown, with small,
punctiform or slightly elongate warts; the outside of the limb is dark green with an obscure
blackish purplish flush and a few scattered, small whitish dots, or brown with a similar
spotting, its inside bright green or brownish with a few small, whitish spots. The spadix
is sessile, shorter than or nearly as long as spathe, up to 17 cm long. The pistillate zone is
cylindric or slightly obconic, up to 1.5 cm long, flowers congested; the ovary is bright pale
green, depressed globose, 2.5–3 mm in diam., 2-loculed; the style is straight, short, 1 mm
long; the stigma is dirty pale yellowish brownish, depressed, large, ca. 2–2.5 mm in diam.,
orbicular or oval in cross section, entire or shallowly or distinctly 2-lobed, lobes rounded.
The staminate zone is slightly or strongly obconic, up to 4 cm long, flowers congested.
Staminate flowers white. The appendix is conic, oval-elliptic, or narrowly fusiform, up to
12 cm long, obtuse, white, with many shallow, often longitudinally confluent depressions
and with or without scattered, tiny, punctiform pustules, base with sterile stamens, gradu-
ally disappearing upwards. Fruits are glossy blue (Figures 2.95 through 2.98).
FIGURE 2.95
Amorphophallus tonkinensis: leaf. (Courtesy of M. Sizemore.)
FIGURE 2.96
Amorphophallus tonkinensis: petiole. (Courtesy of M. Sizemore.)
FIGURE 2.97
Amorphophallus tonkinensis: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.98
Amorphophallus tonkinensis: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
Distribution: China: southeastern Yunnan; northern Vietnam, Laos.

Ecology: In dense tropical forests, moist shaded places; 800–900 m altitude, flowering in May.
NO T E : Amorphophallus tonkinensis is a member of the blue-fruited clade in subg.

Metandrium. It is by far the largest species of this clade and with a two or three other mem-
bers (a. o. A. croatii Hett. & A. Galloway), the only ones to inhabit everwet forests.
2.3.19 Amorphophallus variabilis Blume

In: Blume: Rumphia 1: 146, 1837. t. 35, 37h
The tuber is depressed globose, up to ca. 20 cm in diam., producing numerous annual off-
sets, these shortly spindle-shaped. Leaf solitary with the petiole up to 120 cm long, smooth
or slightly rugose at the base, entirely green (var. immaculatus Hassk.) or a green, olive-green
or dark brown background colour, variously variegated with large and small, confluent and
free, oval, rounded or elongate spots of dark green, white, greyish green, extremely vari-
able. The lamina may be up to 125 cm across, with the leaflets elliptic to elongate elliptic,
acuminate to long acuminate, up to 34 cm long, upper surface moderately glossy, mid green.
Inflorescence solitary, long peduncled; peduncle as petiole, up to 120 cm long. The spathe
is erect, elongate triangular, acute, a slight constriction between base and limb, up to 23 cm
long; the limb near the base suddenly narrowed, margin of limb strongly reflexed, outside
entirely green or dirty green with scattered, small, black dots, to the margin suffused with
brown, the veins pale greenish, inside base maroon, reddish brown, orange or yellowish,
the rest creamy white, limb inside creamy white, pale green or pale reddish brown, base
within with numerous, small, laterally flattened warts and oozing out a clear fluid during
anthesis. Spadix sessile, usually much longer than spathe, up to 58 cm long. The pistillate
zone is cylindric or slightly conic, up to 4 cm long, pistils congested. Ovaries depressed, sub-
circular or diamond-shaped in cross-section, bright green, 3–4 mm in diam., 2- or 3-locular;
style 0.5–2.5 mm long, pale green or purple; stigma oval, elliptic, triangular or subquadran-
gular in cross-section, 1–2.5 mm in diam., yellow, dirty yellow or pale brownish, shallowly
or strongly bilobate, trilobate or quadrilobate. The staminate zone is cylindric or slightly
fusiform or slightly obconic, up to 6 cm long, flowers congested, off-white. The appendix is
narrowly elongate conic, acute, sometimes slightly laterally compressed, up to 48 cm long,
creamy white, greyish brown or pale brown, emitting a powerful gaseous smell, surface
with minute warts and with several shallow, elongate grooves, deepest and most numerous
near the appendix-base. Fruits are orange-red (Figures 2.99 through 2.104).
FIGURE 2.99
Amorphophallus variabilis: tuber with offsets. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.100
Amorphophallus variabilis: petiole detail. (Courtesy of C. Claudel.)
FIGURE 2.101
Amorphophallus variabilis: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.102
Amorphophallus variabilis: spathe and spadix side view. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.103
Amorphophallus variabilis: pistillate and staminate zones. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.104
Amorphophallus variabilis: fruiting. (Courtesy of C. Claudel.)
Distribution: Indonesia: Java, Madura and the Kangean Archipelago, Bali.

Ecology: common everywhere on Java in secondary vegetation with some shade but
never in deep shady forest, also in plantations, in teak forests and in villages, on
many soil types but never on swampy places, 0–700 m altitude. Flowering mostly
June to December, more rarely in other seasons, fruiting from July to January.
NO T E : Amorphophallus variabilis is member of subg. Amorphophallus and sister species

to A. boyceanus Hett (West Malaysia). Local people cook and eat ripe fruits and the peti-
ole (after scraping off the epidermis; Backer, 1914: 11). Leaves are fed to Gourami fishes.
During the first half of this century, tubers of A. variabilis were exported from Java to
Japan, where it was used to prepare Konnyaku-flour. Tubers were also bought by Chinese
tradesmen. In the US, in the late thirties, an interest grew in Ilesmannane-flour, prepared
from Amorphophallus-tubers. Amorphophallus variabilis has repeatedly been reported from
outside Java (e.g., Philippines, Thailand, India, Sri Lanka, Malaysia) but these records are
all based on misidentifications.
2.3.20 Amorphophallus xiei H. Li & Z. L. Dao

In: Novon 16: 240. 2006.
The tuber is dark brown outside, pink inside, depressed globose, up to ca. 16 cm in
diam., rootscars not swollen, without offset development. The leaf appears solitary with
the petiole pink or green at the base, the rest green to dark green, sometimes with some
inconspicuous, paler, rhombic, linear, or irregular pale spots with a dark center or with
scattered very small, dark green dots, 60–80 cm long, glabrous; the lamina is 60–120 cm
across. The leaflets are elliptic, up to ca. 23 cm long, apex acuminate, upper surface mid
green. On the main branching points of the lamina epiphyllar bulbils develop (identi-
cal those in A. bulbifer and A. muelleri: see Section 2.3.4 and 2.3.13). The inflorescence is
short pedunculate, with the peduncle up to 18 cm long. The spathe is almost erect, up
to ca. 18 cm long, outside very pale whitish pinkish with scattered punctiform dark
brown dots and the margin sometimes reddish; inside pale to mid pink, the base often
darker reddish pink, minutely warty. The spadix is sessile, slightly or distinctly longer
than the spathe, 20–24 cm long, The pistillate zone is cylindric, up to 4.5 cm long, flow-
ers congested; ovary pink to reddish pink, obconic, ca. 2 mm in diam., 1-loculed; style
almost absent; stigma nearly sessile, yellow, strongly depressed, disciform, ca. 3 mm
in diam., wider than ovary, slightly 5- or 6-lobed to irregularly 8-lobed. The staminate
zone is cylindric or slightly obconic, up to 7 cm long, staminate flowers congested, pink.
The appendix is off-white or pale pink, fusiform-conic, laterally compressed in larger
individuals, up to 11 cm long, smooth. Fruits are red, developing without pollination
FIGURE 2.105
Amorphophallus xiei: leaf. (Courtesy of C. Claudel.)
FIGURE 2.106
Amorphophallus xiei: lamina with bulbils. (Courtesy of unknown.)
FIGURE 2.107
Amorphophallus xiei: petiole detail. (Courtesy of C. Claudel.)
FIGURE 2.108
Amorphophallus xiei: inflorescence. (Courtesy of H. Li.)
FIGURE 2.109
Amorphophallus xiei: inflorescence. (Courtesy of Y.J. Tao.)
FIGURE 2.110
Amorphophallus xiei: pistillate and staminate zones. (Courtesy of A. Galloway.)
FIGURE 2.111
Amorphophallus xiei: fruiting. (Courtesy of H. Li.)
Distribution: China, western Yunnan (Longchuan).

Ecology: in forest margins, tropical thickets; 900–1,100 m altitude.
NO T E : For comments by the author of the species status of A. xiei, see Li & Hetterscheid,
2010. Amorphophallus xiei shows a mixture in morphological characters of both of its closest
relatives, A. bulbifer and A. muelleri. In terms of molecular phylogenetic relationships, it is
just a bit closer to A. bulbifer (Claudel et al., 2017). These three species form a very interest-
ing triad of apomictic species, with probably all there a triploid genome (established for
A. bulbifer and A. muelleri).
2.3.21 Amorphophallus yuloensis H. Li.

In: J. Wuhan Bot. Res. 6: 211. 1988.
The tuber is depressed globose, up to ca. 10 cm in diam., without offset development.
Petiole surface uniformly olive-green to pale green, up to 75 cm long, smooth; the lamina
up to 100 cm across and on the central branching point and several more distal nodes
intercalary bulbils develop. The leaflets are elliptic to elongate elliptic, up to 25 cm long,
apex acuminate. Inflorescence solitary, short pedunculate; peduncle dark olive-green
with fine, darker, short striations/spots, up to ca. 10 cm long. The spathe is erect, short
convolute, strongly concave, broadly ovate, ca. 11 cm long, margins incurved, obtuse,
outside base pale greenish with small, blackish green dots, center very pale pinkish with
similar dots, distally dirty creamish with scattered, greyish black dots, inside base pale
pink with numerous white verrucae, limb dirty cream. The spadix is subsessile, shorter
than spathe, ca. 8.5 cm long. The pistillate zone cylindric, 1–2 cm long, flowers congested
or slightly separated; ovary large, depressed, orbicular, ca. 4 mm in diam., pale green
with very thin, scattered pinkish, short stripes on the apical half, 2-locular; style short
but distinct; stigma pale dirty yellow, disciform, ca. 2.5 mm in diam., with a shallow
central depression. The staminate zone slightly fusiform, ca. 4 cm long, flowers mostly
separated but distally congested; staminate flower off-white. The appendix is broadly
conic, up to 4 cm long, off-white, glabrous, base irregular, apex obtuse. Fruits are blue
FIGURE 2.112
Amorphophallus yuloensis: tuber. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.113
Amorphophallus yuloensis: leaf bulbil. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.114
Amorphophallus yuloensis: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.115
Amorphophallus yuloensis: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.116
Amorphophallus yuloensis: pistillate zone. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.117
Amorphophallus yuloensis: fruiting. (Courtesy of C. Claudel.)
Distribution: China: Yunnan.

Ecology: In dense primary evergreen valley forests on limestone; 200–2,400 m
altitude.
A rather small species but a clear member of the blue-fruited clade in subg.
NO T E :
Metandrium. Its more detailed phylogenetic position is unclear.
2.3.22 Amorphophallus yunnanensis Engl.

In: Engler, Pflanzenr. IV, 23C (Heft 48): 109. 1911. Synonym: A. kerrii N. E. Br.
The tuber is subglobose or depressed-globose, up to 30 cm in diam., seasonally
developing several offsets, which are rounded or elliptic. The leaf is solitary, with a
smooth petiole of up to 120 cm long, medium to dark olive-green or dark olive-brown

with several rhombic or elliptic-elongate, pale whitish greenish spots. The lamina is
up to 190 cm across, with elliptic leaflets of up to 42 cm length and which are dark
green, often with a bluish sheen when young, margin often narrowly violet. The inflo-
rescence is solitary, long pedunculate; the peduncle may reach 60 cm in length and
looks like the petiole. The spathe is erect, strongly concave, arching over the spa-
dix, broadly ovate, up to 29 cm long, obtuse or acute, base shortly convolute, interior
base smooth or with a few scattered, punctiform warts, exterior white or pale green-
ish white, rarely dark green, sometimes with paler, occasionally ring-like spots near
the base, or flushed pale pinkish, the margin sometimes lined pinkish, interior pale
greenish white without spots, limb exterior dirty creamy, sometimes with faint spot-
ting, margin sometimes pale pinkish violet, interior creamy or pale greenish white,
the margin sometimes pale pinkish violet. The spadix is much shorter than spathe,
up to 18 cm long, stipitate; the pistillate zone is cylindrical, slightly conical or obconi-
cal, up to 3.5 cm long, flowers congested though sometimes loosely arranged in the
lower part. The ovaries are globose, subglobose or depressed, circular or angular in
outline, 2–4 mm in diam., green, pale green, brownish green or purplish; the style
is slender to shortly conical, straight or curved; the stigma is variable, usually dis-
tinctly broader than the style diameter, disciform to subhemispherical, more rarely
superficial, punctiform, circular or oval in outline, with a shallow central depression
to clearly bilobed, pale yellowish or dirty brownish. The staminate zone is conical
or fusiform-c ylindrical, rarely obconical, sometimes (partly) laterally compressed, up
to 4 cm long, flowers congested; staminate flowers, creamy white or partly maroon
or dark brown. The appendix is broadly ovate, conical or triangular-ovate, rarely
subcylindrical, up to 11 cm long, inflated, or strongly laterally compressed, obtuse,
rarely acute, creamy white or pale pinkish, base strongly truncate, usually with a
few broadly conical staminodes, surface smooth or verruculate, rarely entirely echi-
nate, often with a few longitudinal, shallow or deep, broad folds, or irregularly folded
throughout. Fruits are blue or violet (Figures 2.118 through 2.125).
FIGURE 2.118
Amorphophallus yunnanensis: tuber. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.119
Amorphophallus yunnanensis: petiole detail. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.120
Amorphophallus yunnanensis: petiole detail. (Courtesy of C. Claudel.)
FIGURE 2.121
Amorphophallus yunnanensis: inflorescence. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.122
Amorphophallus yunnanensis: inflorescence, lateral view. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.123
Amorphophallus yunnanensis: spadix exposed. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.124
Amorphophallus yunnanensis: pistillate zone. (Courtesy of W.L.A. Hetterscheid.)
FIGURE 2.125
Amorphophallus yunnanensis: fruits maturing to blue. (Courtesy of C. Claudel.)
Distribution: Thailand: Mae Hong Son, Chiang Mai, Lamphun, Tak, Phitsanulok,
Khon Kaen, Chaiyaphum (Phu Khieo); China: Guangxi, Guizhou, Yunnan; Laos;
northern Vietnam.
Ecology: in shaded places in primary evergreen or mixed evergreen/deciduous for-
ests, on metamorphic bedrock, in rich soils, or secondary forests, thickets, forest
margins; 100–3,300 m altitude.
NO T E S :Amorphophallus yunnanensis is again a typical member of the blue-fruited clade in

subg. Metandrium. It is very similar in appearance to A. kachinensis and A. corrugatus. Both
species, however, have a unilocular ovary and an often deeply fissured, stipitate appendix,
with the spathe mottled with reddish purple spots, especially on the inner side.
2.4 Growth Cycles

Most Amorphophallus species occur in areas with seasonality in either temperature, pre-
cipitation or both. Most of these species therefore exhibit a short or prolonged dormancy
period during which only the underground storage (tuber) organ remains. In immature
specimens growth after dormancy starts every time with leaf development but when a
plant reaches flowering size dormancy may be followed by the development of a solitary
inflorescence. After pollination and fruit set a new dormancy period sets in, after which
in the next season a leaf will develop. This pattern repeats itself. It may be that in case
of failed pollination, a leaf may develop after all. This often happens in several species
in cultivation but there are no observations like this in natural circumstances. During
a leaf phase, the tuber is depleted and a full new tuber develops, which is usually two
or more times larger than the previous one. This tuber develops a terminal bud, from
which in the next season the inflorescence will develop at maturity. During flowering
and fruiting, a part of the energy and mass of the tuber is used. The leaf to follow,
develops from a bud, laterally placed from the terminal scar, left after the peduncle has
rotted off. This cycle type is common to most of the species treated in this book. Notable
exceptions are A. tonkinensis and A. coaetaneus. The former grows in everwet forests and
shows no strong seasonality, often developing a leaf during fruit set or shortly after the
fruiting parts have been eaten and rotted off. Amorphophallus coaetaneus is an evergreen
species and has no clear leafless dormancy. Both species represent a different cycle type
each, shared by a few (in case of A. coaetaneus, all developing rhizomes) or several spe-
cies (in case of A. tonkinensis, several other species dwelling in everwet forests). They will
not be treated here but await a publication by the author on morphological evolution in
Amorphophallus.
2.5 Glucomannan Content in Various Amorphophallus Species

The Amorphophallus species described in Section 2.3 store their reserve polysaccharides,
starch and glucomannan, in underground tubers.
In addition, there are species that contain small amounts of alkaloids and toxic sub-
stances such as oxalic acid in fresh tubers. Since it is difficult to remove those undesir-
able substances during processing, such species have no commercial value for the food
industry. However, some of these species contain considerable amounts of konjac gluco-
mannan (KGM). This polymer is composed of D-glucose and D-mannose connected by
β-1,4 glycosidic bonds in a molecular ratio of 1:1.6 to 4.2 (Chen et al., 2005). KGM can have
high viscosity at low concentration and can therefore be used as a raw material in many
industries (Sheng and Teng, 2008). Amorphophallus is the main plant genus used for com-
mercial production of glucomannan. The glucomannan content in tubers of some species
is approximately 60% of their dry weight (Diao et al., 2014). This is why the glucomannan
content is an important quality indicator.
The size of the tubers increases during the growing season and it may reach up to three
times its former weight per season (Zhao, 2010). An approximately three-year-old tuber is
suitable for KGM flour production. Most of the species with high glucomannan content are
grown in plantations in China and Japan. They are also grown in Southeast Asia but at a
smaller scale (Zhou et al., 2004).
The species with high KGM content are as follows:
Amorphophallus konjac K. Koch: an important crop cultivated in Japan and China for
the KGM flour production industry. It is a diploid species (2n = 26). The tuber con-
sists of 49%–60% (w/w) water soluble KGM (Li et al., 2005).
A. albus is a native Chinese species and mainly cultivated as crop in Yunnan, south-
west China (Long, 1998). It is diploid (2n = 26). The tubers of this species contain
around 59.3% KGM (w/w) (Liu, 2004).
A. bulbifer is a triploid (2n = 39) with that produces nucellar, clonal seedlings without
pollination. This species tolerates a wide range of temperatures and grows well
between 15°C and 30°C. A. bulbifer has been successfully introduced and is cul-
tivated in Yunnan, China. The KGM content in this species is between 48%–52%
which is lower than A. konjac and A. muelleri (Zhang et al., 2009, 2010; Zhao, 2010).
It was used as a model for the glucomanan biosynthesis pathway (Diao et al., 2014)
and for a complete chloroplast genome study (Hu et al., 2008).
A. muelleri is also triploid (3n) with 39 chromosomes and is an important wild and
cultivated species in Thailand and Indonesia (Java). It has a high KGM content
(72%–78%) (Zhang et al., 2010; Zhao, 2010). There is a study that reported about a
relationships between phylogeny and glucomanan content of Amorphophallus spp.
collected in Thailand (Mekkerdchoo et al., 2016). The highest KGM content was
found in A. muelleri (60.16%–68.93%) followed by A. krausei, A. kachinensis, A. bulbifer,
A. xiei and A. corrugatus respectively. Moreover, the study retrieved two monophy-
letic clades using DNA sequencing with high KGM content species (Figure 2.126).
The RAPD analysis generated specific bands identifying species belonging only to
the high and medium glucomamnan content clades. It can be used as a screening
tool for economic Amorphophallus species. Moreover, these species can also be stud-
ied for complete chloroplast genome construction (Hu et al., 2008) and ceramidase
production from sphingolipid metabolic pathway (Zhong et al., 2018).
The glucomannan content can be affected by various factors such as location, soil, weather,
age of tuber and processing (Fang and Wu, 2004; Zhang and Liu, 2006; Zhang et al., 2005).
Liu (2004) found that the KGM content of A. konjac grown in different areas varied slightly
FIGURE 2.126
Phylogenetic tree of Amorphophallus spp. in Thailand and glucomannan content. (From Mekkerdchoo, O. et al.,
Phytotaxa, 282, 81–106, 2016.)
(ranging from 58.8% to 52.1%). However, the results of these studies also indicated that the
individual species is the main factor determining the KGM content and the quality of the
KGM flour and that the growing conditions are a secondary factor that can be optimized
to maximize the KGM yield.
2.6 Hybrids
2.6.1 Hybridization in Amorphophallus
During the phylogenetic investigations of the genus Amorphophallus on a larger scale
(Claudel et al., 2017), a negligible but intriguing phenomenon constantly reproduced in
most analyses: four species of the subg. Scutandrium Hett. & Claudel paired in an unex-
pected combination. The first two species, A. konjac K. Koch and A. maxwellii Hett., are tall
and large. The plants easily exceed one meter height and are characterized by an equally
large inflorescence, both with a dark maroon spathe (Hetterscheid and Ittenbach, 1996).
In contrast, the other two species which are ‘morphologically very similar’ (Hetterscheid
and Ittenbach, 1996) are distinctly smaller and the inflorescences bear light-coloured
spathes. The spathe is whitish to pale green outside and faintly maroonish inside in the
case of A. krausei Engl; and greenish outside and whitish inside in the case of A. albus
P. Y. Liu & J. F. Chen. Based on the morphological similarity between the two species pairs,
A. konjac was expected to be closely related to A. maxwellii and A. albus was expected to
pair with A. krausei. Instead, A. konjac paired with A. albus and A. maxwellii with A. krausei.
This result was too intriguing to be ignored and, as a consequence different accessions of
A. albus and A. konjac were sequenced in order to validate or disprove the result. However,
it persisted and the question arose if hybridization events possibly influenced the result.
It first seemed unlikely, however, taking into consideration that several Amorphophallus
species, (i.e., A. paeoniifolius (Dennst.) Nicolson, A. albus and A. konjac) have a long breed-
ing history in Asia (Hetterscheid and Ittenbach, 1996; Zheng et al., 2013; Liu et al., 2015) it
did not seem impossible. Moreover, Zhang et al. (1998) reported the successful hybridiza-
tion of several Amorphophallus species, amongst others hybridization between A. albus and
A. konjac. It therefore seemed suddenly not only possible but even likely that these species
might have been hybridized in the past.
Thus, the first author (Claudel et al., 2017) decided to reproduce the cross between the
aforementioned species in order to investigate the placement of the progeny within the
phylogenetic analysis. This was the starting point of further attempts to cross as many dif-
ferent Amorphophallus species as possible and to explore the limits of hybridization within
the genus. Thanks to Mr. John Tan, an avid plantsman from Singapore, the author received
the necessary support to raise many hundred plants of hybrid origin. Moreover, several
Amorphophallus enthusiasts around the world, from Australia to Indonesia, from the US
to Europe joined the informal project and started to hybridize species and consequently
reported the outcome. The overall aim of the project was to create new hybrids of orna-
mental value, with desirable traits, such as increased hardiness and robustness and/or
decreased odour properties (Claudel and Galloway, 2012). Therefore, the limits of hybrid-
ization within the genus had to be explored in order to improve or to combine specific
traits from different species and to increase the overall vigour.
Although the ornamental value was in the foreground instead of crop traits, such as
tuber growth or glucomannan content, a short comment should be made concerning
species with agricultural importance. These essentially include six species from three
different subgenera, namely A. albus and A. konjac, A. paeoniifolius and A. bulbifer Blume,
A. muelleri Blume and A. xiei H. Li & Z. L. Dao.
Amorphophallus albus and A. konjac belong to the subgenus Scutandrium and both have
2n = 26 chromosomes which is considered to be the basic chromosome number in the
genus (Shete et al., 2015). As previously mentioned these two species are compatible and
the hybrids show signs of hybrid vigour (Claudel et al., 2013). Moreover, the resulting
F1-generation is fertile and the hybrids can either be crossed one with each other or back-
crossed with one of the parents (personal observation). This opens up endless opportuni-
ties for breeding programs of suitable crop plants. Overcoming the poor disease resistance
of A. konjac (Zheng et al., 2013) could be a possible outcome. Moreover, the study from
Zheng et al. (2013) concerning A. konjac revealed that the genetic ‘variation between wild
and cultivar genotypes’ is very small. Yin et al. (2019) come to the same conclusion concern-
ing the genetic variation within A. albus. In other words, it seems that the Amorphophallus
species have a long history of cultivation but not necessarily a long breeding history.
Amorphophallus paeoniifolius from the subgenus Amorphophallus is the species with
the widest distribution, possibly due to their use as starchy crop plant (Hetterscheid &
Ittenbach, 1996). It belongs to a small clade (Claudel et al., 2017) together with A. prainii Hook
f. and a few other species which are characterized by 2n = 28 chromosomes (Chauhan &
Brandham, 1985). Although widely planted, the aberrant chromosome number limits the
potential in terms of hybridization as only few species can be used as crossing partners.
Even more limited in this context are the three species A. bulbifer, A. muelleri and A. xiei
from the subgenus Metandrium Stapf. The chromosome number for A. xiei has not been
investigated; however, A. xiei is so similar to A. bulbifer that its species identity has been
questioned by Li and Hetterscheid (2010). Amorphophallus bulbifer and A. muelleri are trip-
loids characterized by the chromosome number 2n = 39 (Ramachandran, 1977; Chauhan
and Brandham, 1985; Patil, 1995; Shete et al., 2015) and it has been discussed if the chromo-
some number in these two species is based on allotriploidy or autoploidy (Chauhan and
Brandham, 1985; Patil, 1995; Shete et al., 2015). Both species seemingly exclusively rely on
the formation of epiphyllar bulbils and apomictic seed formation for propagation (more
precisely: agamospermy). This is underlined by the investigation of Patil (1995) who states
that the: ‘autopolyploid nature in A. bulbifer (2n = 39) is supported by the lowest pollen fer-
tility 5.69% and apomictic seed formation...’ Thus, these species seemingly cannot be used
for generative propagation, neither for hybridization nor for selections based on breeding.
However, Kuruvilla et al. (1989) reported 2n = 26 for A. bulbifer. Possibly some popula-
tions from A. bulbifer and A. muelleri are true diploids which could be used for breeding
experiments. This finding would fit with the concept that apomixis in plants is facultative
(Savidan, 2000), in other words that if sexual reproduction has not been observed, than the
observation has just not been rigorous enough. Moreover, although the pollen fertility is
very low (Patil, 1995) the pollen nevertheless is fertile. These points deserve closer atten-
tion as these species are of economic value (Zhao et al., 2009, 2010; Zhang et al., 2010) and
breeding programs could give rise to new high-performance cultivars. It is noteworthy to
add that Tjio (1948) reports 2n = 39 for A. rivieri Dur., a synonym of A. konjac. Either the
plant was not correctly identified and was in fact a specimen of A. bulbifer or A. muelleri, or
further triploid specimens occur within other species.
However, at the starting point the focus was set on the ornamental value and the fea-
sibility of a cross. As a result, nearly 250 crosses worldwide were performed, of which
47 were successful and yielded viable seeds (Claudel and Galloway, 2012; Claudel
et al., 2013; Claudel and Mangelsdorff, 2014). Some outstanding specimens from a few
crosses were named and subsequently released, namely: Amorphophallus ‘Kiat Tan’
(A. lewallei Malaisse & Bamps x A. maximus (Engl.)), Amorphophallus ‘Mary Sizemore’
and Amorphophallus ‘Meister Eckhardt’ (both A. albus x A. konjac), Amorphophallus ‘Heine’
(A. lewallei x A. richardsiae Ittenb.), Amorphophallus ‘Blue Nightspot’ (A. glaucophyllus Hett. &
Serebr. x A. lacourii Linden & Andre) and Amorphophallus ‘Majda’ (A. pulchellus Hett. &
Schuit. x A. myosuroides Hett. & A. Galloway) (Claudel, 2019; Galloway, 2019). Besides
exhibiting good growing properties and beautiful foliage or inflorescences it was required
that these could not be mistaken for the true species. These cultivars represent a trial bal-
loon and will demonstrate if Amorphophallus cultivars of hybrid origin will find their niche
and spread in collections.
The perhaps most amazing cross was performed by Ralph Mangelsdorff in 2002
(Claudel et al., 2012). It involved two very different crossing partners in terms of absolute
size. The seed parent was A. variabilis Bl. which usually hardly exceeds 1.20 meter height
(Hetterscheid and Ittenbach, 1996) and has a long peduncled but otherwise rather small
and inconspicuous inflorescence. The pollen parent however was A. titanum (Becc.) Becc.
ex Arcangeli, the most striking species of the genus with leaves up to six meters high and
sessile, very large inflorescences exceeding 3 meter height (McPherson and Hetterscheid,
2011). Only a few seeds fully developed (Claudel et al., 2012) and only one seedling reached
flowering stage. The plant displayed perfectly intermediate character traits and was very
showy at that. It was named Amorphophallus ‘John Tan’ (Claudel et al., 2012) in honour of
Mr. Tan and represents therefore the first named Amorphophallus cultivar of hybrid origin.
Moreover, used as pollen parent, it also is the ‘father’ of the first Amorphophallus hybrid
involving three Amorphophallus species (Claudel et al., 2012).
Since 2014 the interest in hybridization amongst enthusiasts seemingly declined.
Considering that one cross can yield a few to a few hundred seeds and that it takes,
species-dependant, three to seven years to raise the hybrids until maturity, it becomes
apparent that it is a task which requires time, space and efforts, especially if larger species
are involved. Moreover, it requires close observation of the plants, as Amorphophallus spe-
cies, like all aroids are protogynous (Boyce and Wong, 2012) which in most cases imply a
short time frame for successful pollination. As a general rule, the ‘female phase’, the phase
characterized by receptive stigmas, starts and ends on day one of anthesis, whereas the
‘male phase’, characterized by pollen release, occurs on day two. That said, the female
phase does not necessarily last the whole day but can end after a period of six hours only
(personal observation). In contrast to that, some species are characterized by an extended
female phase which lasts up to five days (personal observation) such as for example
A. antsingyensis Bogner, Hett. & Ittenb., A. gigas Teijsm. & Binnend., A. henryi N. E. Br.,
A. konjac, A. lambii Mayo & Widjaja, A. natolii Hett. et al. and A. variabilis. Either way, the
pollen needs to be applied to the stigmas when these are receptive which is often indicated
by a sticky fluid on the stigma surface. Applying pollen either requires a plant which flow-
ered in the preceding days and serves as pollen donator or stored pollen, dried and frozen
or refrigerated (Claudel & Galloway, 2012).
Fresh pollen is naturally the best choice and is ideally applied on the day of release as
it is assumed to be short-lived and to deteriorate within a few days at room temperature
(Harrington, 1970; Zhang et al., 1998; Barabé et al., 2008). However, it can be gently dried
using silica gel (Claudel and Galloway, 2012) and either be refrigerated at 4°C–8°C or frozen
at −20°C or comparable temperatures. As stated in Claudel et al. (2013) and Claudel and
Mangelsdorff (2014), eight out of 25 successful crosses were performed using frozen pollen;
in one case the pollen had been stored for more than two years. Moreover, in the new series
of crosses (see Appendix 2.I) 16 out of 37 successful crosses have been performed using
either refrigerated or frozen pollen. This demonstrates the usefulness of refrigerated/

frozen pollen and it shows the potential of a pollen bank. A pollen bank would not only
ease the task in hybridization attempts but even more in ex-situ conservation projects as
it would allow, for example, self-pollination of single specimens using stored pollen from
former inflorescences. However, except for the fact that Amorphophallus pollen is storable
under the aforementioned conditions, many questions remain to be addressed and inves-
tigated. For example the impact of cooling versus freezing on the longevity of the pollen.
Another area to be investigated is the decrease of viability over time under different cool-
ing or freezing regimes. Furthermore, the effects of further freezing regimes including
−80°C and cryogenic cooling on viability also need to be investigated. Lastly, the question
whether the different kinds of pollen (Van der Ham et al., 1998, 2005; Ulrich et al., 2017)
within the genus Amorphophallus respond differently to the suggested or other treatments
should also be addressed.
It is a fortunate and interesting development that the Toronto Zoo (Ontario, Canada) has
started to address these questions (personal communication, Paul Gellatly, curatorial gar-
dener). Although a zoo, the Toronto Zoo also maintains a growing Amorphophallus collec-
tion, including a dozen specimens of A. titanum. Fortunately, the Toronto Zoo has a strong
focus on conservation genetics and ex-situ conservation projects and houses the facilities
to properly store genetic material. In 2018 the first specimen of A. titanum flowered and
aroused considerable attention. It was decided to collect the pollen for conservation pur-
poses, namely pollinating future A. titanum inflorescences. The pollen has been collected
and stored in various ways to determine the viability of long term pollen storage. It has
been frozen and stored in −80°C, as well as cryogenically preserved. The Zoo is currently
in the process of viability testing in order to determine if these approaches constitute a
practicable long term pollen storage solution.
Despite the work it entails, nearly 250 further pollination attempts (see Appendices 2.I
and 2.II) have been performed in the meantime by the most arduous hybridizers, namely
Alan Galloway from the US and Steve Jackson from Australia. It is worthwhile pointing
out that it was A. Galloway who introduced many Amorphophallus species to science over
the last two decades. Moreover, he is a most skilled grower. Both facts combined have
opened up many hybridization options.
On the other side of the globe, Steve Jackson has created the first hybrids involving spe-
cies such as A. decus-silvae Backer & Alderw. and A. gigas, two of the tallest and most
spectacular species of the genus. The cultivation of these species is not exactly easy and
although S. Jackson has been supported by a favourable climate, the fact that he kept these
species in cultivation for decades speaks for itself.
Except for five pollinations performed by the first author and not taking into account
pollinations which involved several attempts, 229 further hybridizations attempts were
carried out by A. Galloway and S. Jackson. Out of a total of 234 hybridizations attempts
37 were successful and yielded viable seeds (see Appendix 2.I). However, the overall fitness
of the progeny can strongly differ. In general, two categories can be observed. The first one
includes hybrids showing signs of hybrid vigour acquired through heterosis. Heterosis
can be defined as ‘The physiological vigour of an organism as manifested in its rapidity
of growth, its height and general robustness, is positively correlated with the degree of
dissimilarity in the gametes by whose union the organism was formed …’ (Shull, 1948).
In other words, the hybridogenic Amorphophallus progeny is more robust and produces
larger leaves and inflorescences than its corresponding parents. The second category
is the outbreeding depression (not to be confounded with the inbreeding depression).
The hybrids display a loss of fitness, are more vulnerable to diseases and disturbances and
have weaker growing and flowering properties than their corresponding parents (Leimu
and Fischer, 2010; Barmentlo et al., 2018). This is due to biochemical and/or physiological
incompatibilities between the crossing partners. The crossing partners are genetically too
distant and the selective advantage of adapted gene complexes of the involved species
is disrupted through hybridization (Wikipedia, 2019). One of the markers, indicating the
limitation of hybridization in Amorphophallus is albinism (Claudel et al., 2013). Albinism
in the progeny of a given Amorphophallus cross can be displayed at two levels. One is the
number of seedlings affected and the second is the degree of albinism that a single seed-
ling expresses. In the best case only a few seedlings show slight signs of chlorophyll-free
leaf tissue which may eventually slowly turn green after the leaf unfolds. In the worst case
all the seedlings are completely devoid of chlorophyll and will die as soon as the stored
nutrients are consumed.
This raises the question how closely related two crossing partners need to be if ‘optimal
outcrossing distance’ (Schierup and Christiansen, 1996) or hybrid vigour is the goal or
on the opposite, how distantly related they can possibly be if not improvement but pure
survival at any cost is the aim. Unfortunately there is no specific answer to this question.
It must be taken into consideration that effects based on inbreeding depression, outbreed-
ing depression or heterosis play a role even between different populations of a given single
species (Leimu and Fischer, 2010; Barmentlo et al., 2018). Predicting which effect might
dominate would require a precise knowledge about the genetics of each crossing partner.
Besides this limitation it must be also taken into consideration that the presented hybrid-
ization attempts are arbitrary, mainly for two reasons. Firstly, the involved species are
selected based on traits judged desirable by the hybridizers. Second, the opportunities to
hybridize depend on the cultivated species.
However, some concluding observations can be made. Out of nearly 500 hybridization
attempts a total of 84 yielded viable seeds, especially crosses between closely related species
are often successful. As insignificant as this might seem, it demonstrates that Amorphophallus
species hybridize readily which suggests that interspecific hybridization barriers are not pro-
nounced in many species. This could be accounted for by adaptive radiation. Amorphophallus
is a comparatively young genus (Nauheimer et al., 2012) within the Araceae. However, with
an estimated 219 species (Boyce and Croat, 2011) it exhibits a high species diversity growing
in the (sub)tropical zones of the palaeotropics, outranking all other aroid genera in mor-
phological diversity (Hetterscheid and Ittenbach,1996; Claudel et al., 2017). For example, the
genus encompasses an exceptionally high diversity in berry colour, ranging from white,
green, and yellow, orange, red to blue and purple. This is a characteristic trait indicating seed
dispersal through birds (Claudel et al., 2017) which might have played a major role in the
wide distribution of the genus. Last but not least, many closely related species have similar
or even identical nuclear and plastid sequences (Claudel et al., 2017). Although speculative,
all facts combined – high species and morphological diversity acquired in a short period
of time, palaeotropical distribution pattern, birds as dispersal vector and finally, similar
genetic sequences – suggest adaptive radiation.
Additionally, six out of the 37 successful crosses involve three different species and
one cross even involves four species (see Appendix 2.II). The latter consists of a hybrid
between two Malagasy species as pollen acceptor (A. taurostigma x A. ankarana) crossed
with a hybrid between two African species as pollen donor (A. lewallei x A. impressus).
Nine of the successful crosses even involve species from different subgenera. For example
A. variabilis crossed with A. maximus involves A. variabilis, a species from South East Asia
from the subgenus Amorphophallus and A. maximus an African species from the subgenus
Afrophallus Hett. & Claudel.
Simply put, the boundaries of hybridization within Amorphophallus are not yet
reached, there is still a lot of potential to be uncovered and many questions remain to
be answered.
Acknowledgements
The author of Section 2.6 Hybrids, C. Claudel, wants to express his gratitude towards
Mr. John Tan from Singapore who made the hybrid possible from the beginning to the
end. Furthermore, the author wants to thank Steve Jackson, Alan Galloway and Bjoern
Malkmus-Hussein for joining the journey.
Appendix 2.I Successful Amorphophallus Hybridizations
Pollen Acceptor Pollen Donor Pollen Used Hybridizer Result

(A. decus-silvae Backer & (A. decus-silvae Refrigerated S. Jackson Success
Alderw. x A. variabilis Bl.) Backer & Alderw. x
A. variabilis Bl.)
(A. lewallei Malaisse & A. henryi N. E. Br. Refrigerated S. Jackson Success, three species
Bamps x A. impressus + subgeneric cross
Ittenb.)
(A. lewallei Malaisse & (A. impressus Ittenb. x Fresh S. Jackson Success, three species
Bamps x A. impressus A. taurostigma
Ittenb.) Ittenb. & Hett ‘White
Veins’)
(A. taurostigma Ittenb. & (A. lewallei Malaisse & Fresh S. Jackson Success, four species
Hett x A. ankarana Hett.) Bamps x A. impressus
Ittenb.)
(A. variabilis Bl. x A. longituberosus (Engl.) Fresh S. Jackson Success, three species
A. decus-silvae Backer & Engl. & Gehrm. + subgeneric cross
Alderw.)
A. decus-silvae Backer & A. borneensis (Engl.) Refrigerated S. Jackson Success
Alderw. Engl. & Gehrm.
A. decus-silvae Backer & A. henryi N. E. Br. Fresh S. Jackson Success, subgeneric
Alderw. cross
A. discophorus Backer & A. galbra F.M. Bailey Refrigerated S. Jackson Success
Alderw.
A. galbra F.M. Bailey (A. variabilis Bl. x Fresh S. Jackson Success
A. decus-silvae
Backer & Alderw.)
A. henryi N. E. Br. (A. lewallei Malaisse & Fresh S. Jackson Success, three species
Ittenb.)
A. henryi N. E. Br. A. laoticus Hett. Fresh S. Jackson Success
A. hewittii Alderw. (A. decus-silvae Fresh S. Jackson Success, three species
Backer & Alderw. x
A. variabilis Bl.)
(Continued)

A. impressus Ittenb. A. maximus (Engl.) Refrigerated A. Galloway Success
AGA-1158-01 N. E. Br. AGA-0240-01
A. konjac K. Koch A. albus P.Y. Liu & J.F. Frozen A. Galloway Success
AGA-1266-01 Chen
A. konjac K. Koch A. albispathus Hett. Frozen A. Galloway Success
AGA-1797-01
A. konjac K. Koch A. crispifolius Frozen A. Galloway Success
AGA-1798-01 A. Galloway,
A. Ongsakul, &
P. Schmidt
A. konjac K. Koch A. kachinensis Engl. & Fresh A. Galloway Success
AGA-1798-01 Gehrm. AGA-2500-01
A. laoticus Hett. (A. lewallei Malaisse & Refrigerated S. Jackson Success, three species
Ittenb.)
A. lewallei Malaisse & A. antsingyensis Bogner, Fresh C. Claudel Success
Bamps Hett. & Ittenb.
A. lewallei Malaisse & A. gomboczianus Pic. Fresh C. Claudel Success
Bamps Serm.
A. longiconnectivus Bogn. A. schmidtiae Hett. & Frozen A. Galloway Success
AGA-0891-01 A. Galloway
AGA-2188-01
A. maximus (Engl.) N. E. Br. A. variabilis Bl. Fresh S. Jackson Success, subgeneric
cross
A. natolii Hett.et al. A. variabilis Bl. Refrigerated S. Jackson Success, subgeneric
cross
A. ochroleucus Hett. & V.D. A. dunnii Tutch Fresh S. Jackson Success
Nguyen
A. ongsakulii Hett. & A. A. myosuroides Hett. & Fresh A. Galloway Success
Galloway AGA-1534-01 A. Galloway
AGA-1756-01
A. operculatus (ined.) A. sizemoreae Hett. Refrigerated S. Jackson Success
A. pulchellus Hett. & Schuit. A. ongsakulii Hett. & Fresh A. Galloway Success
AGA-2342-01 A. Galloway
A. richardsiae Ittenb. A. mossambicensis Refrigerated A. Galloway Success
AGA-1920-01 (Schott ex Garcke)
N. E. Br. AGA-0900-01
A. thaiensis S.-Y. Hu A. putii Gagn. Fresh A. Galloway Success
AGA-2236-01 AGA-0832-01
A. variabilis Bl. A. natolii Hett. et al. Frozen S. Jackson Success (x2),
subgeneric cross
A. variabilis Bl. (A. variabilis Bl. x Fresh S. Jackson Success
A. decus-silvae
Backer & Alderw.)
A. variabilis Bl. A. lambii Mayo & Refrigerated S. Jackson Success
Widjaja
A. variabilis Bl. A. maximus (Engl.) Fresh S. Jackson Success
N. E. Br.
A. variabilis Bl. ‘Slukas (A. variabilis Bl. x Fresh S. Jackson Success
Dark Giant’ A. decus-silvae
Backer & Alderw.)
(Continued)

A. variabilis Bl. ‘Slukas A. dracontioides (Engl.) Frozen S. Jackson Success
Dark Giant’ N. E. Br.
A. variabilis Bl. ‘Slukas A. hewittii Alderw. Fresh S. Jackson Success
Dark Giant’
A. yunnanensis Engl. A. dunnii Tutch. Fresh S. Jackson Success
Appendix 2.II Failed Amorphophallus Hybridizations
Pollen Acceptor Pollen Donor Pollen Hybridizer Result

(A. decus-silvae Backer & A. henryi N. E. Br. Refrigerated S. Jackson Failed (x2)
Alderw. x A. variabilis Bl.)
(A. decus-silvae Backer & A. dracontioides (Engl.) N. E. Br. Frozen S. Jackson Failed
(A. decus-silvae Backer & A. laoticus Hett. Refrigerated S. Jackson Failed
(A. decus-silvae Backer & A. longituberosus (Engl.) Frozen S. Jackson Failed
Alderw. x A. variabilis Bl.) Engl. & Gehrm.
(A. decus-silvae Backer & A. taurostigma Ittenb. & Hett Fresh S. Jackson Failed
Alderw. x A. variabilis Bl.) ‘White Veins’
(A. decus-silvae Backer & A. laoticus Hett. Frozen S. Jackson Failed
(A. excentricus Hett. x A. scutatus Hett. & T.C. Refrigerated S. Jackson Failed
A. krausei Engl.) Chapman
(A. impressus Ittenb. x A. elatus Ridl. Frozen S. Jackson Failed
A. taurostigma ‘White Veins’)
(A. lewallei Malaisse & Bamps A. rostratus Hett. Fresh S. Jackson Failed
x A. impressus Ittenb.)
(A. lewallei Malaisse & Bamps A. laoticus Hett. Fresh S. Jackson Failed
(A. maximus (Engl.) N. E. Br. x A. natolii Hett.et al. Refrigerated S. Jackson Failed
A. variabilis Bl.)
(A. ongsakulii Hett. & A. titanum (Becc.) Becc. ex Frozen A. Galloway Failed
A. Galloway x A. myosuroides Arcangeli
Hett. & A. Galloway
(A. variabilis Bl. x A. borneensis A. maxwellii Hett. Frozen A. Galloway Failed
(Engl.) Engl. & Gehrm.)
AGA-2729-09
(A. variabilis Bl. x A. decus- A. laoticus Hett. Refrigerated S. Jackson Failed
silvae Backer & Alderw.)
(A. variabilis Bl. x A. decus- A. natolii Hett. et al. Frozen S. Jackson Failed (x2)
(A. variabilis Bl. x A. decus- (A. lewallei Malaisse & Bamps Frozen S. Jackson Failed
silvae Backer & Alderw.) x A. impressus Ittenb.)
(A. variabilis Bl. x A. decus- A. discophorus Backer & Refrigerated S. Jackson Failed
silvae Backer & Alderw.) Alderw.
(A. variabilis Bl. x A. decus- A. rostratus Hett. Refrigerated S. Jackson Failed
(Continued)

(A. variabilis Bl. x A. decus- A. taurostigma Ittenb. & Hett Refrigerated S. Jackson Failed
silvae Backer & Alderw.) ‘White Veins’
A. albus P.Y.Liu & J.F.Chen A. cirrifer Stapf Refrigerated A. Galloway Failed
A. albus P.Y. Liu & J.F. Chen A. maxwellii Hett. Refrigerated A. Galloway Failed
A. albus P.Y. Liu & J.F. Chen A. natolii Hett. et al. Frozen S. Jackson Failed
A. amygdaloides Hett. & M. A. brevipetiolatus A. Galloway, Frozen A. Galloway Failed
Sizemore AGA-1047-03 A. Ongsakul, & P. Schmidt
AGA-1570-03
A. amygdaloides Hett. & M. A. muelleri Bl. Fresh A. Galloway Failed
Sizemore AGA-1047-03
A. asterostigmatus Bogn. & (A. decus-silvae Backer & Frozen A. Galloway Failed
Hett. AGA-1964-01 Alderw. x A. variabilis Bl.)
AGA-2461-11
A. atrorubens Hett. & M. A. maxwellii Hett. Refrigerated A. Galloway Failed
Sizemore AGA-1214-01
A. atroviridis Hett. A. maxwellii Hett. Refrigerated A. Galloway Failed
AGA-1046-01 AGA-1177-01
A. bangkokensis Gagn. A. rostratus Hett. Refrigerated S. Jackson Failed
A. barbatus A. Galloway & A. laoticus Hett. Fresh C. Claudel Failed
A. Ongsakul
A. barbatus A. Galloway & A. titanum (Becc.) Becc. ex Frozen A. Galloway Failed
A. Ongsakul AGA-2309-01 Arcangeli
A. borneensis (Engl.) (A. decus-silvae Backer & Refrigerated S. Jackson Failed
Engl. & Gehrm. Alderw. x A. variabilis Bl.)
A. borneensis (Engl.) A. interruptus Engl. & Gehrm. Refrigerated S. Jackson Failed
Engl. & Gehrm.
A. borneensis (Engl.) A. titanum (Becc.) Becc. ex Frozen S. Jackson Failed
Engl. & Gehrm. Arcangeli
A. boyceanus Hett. A. gomboczianus Pic. Serm. Frozen S. Jackson Failed
A. boyceanus Hett. A. scutatus Hett. & T.C. Refrigerated S. Jackson Failed
Chapman
A. boyceanus Hett. A. tuberculatus Hett. & V.D. Refrigerated S. Jackson Failed
Nguyen.
A. cicatricifer Ridl. (A. lewallei Malaisse & Bamps Refrigerated S. Jackson Failed
A. cicatricifer Ridl. (A. lewallei Malaisse & Bamps Refrigerated S. Jackson Failed
A. cicatricifer Ridl. A. sizemoreae Hett. Fresh S. Jackson Failed
A. coaetaneus Liu & Wie (A. excentricus Hett. x Frozen S. Jackson Failed
A. krausei Engl.)
A. coaetaneus Liu & Wie A. kachinensis Engl. & Gehrm. Frozen A. Galloway Failed
AGA-1800-01 AGA-1815-01
A. crispifolius A. Galloway, A. laoticus Hett. AGA-2025-01 Refrigerated A. Galloway Failed
A. Ongsakul, & P. Schmidt
AGA-1753-01
A. croatii Hett. & A. Galloway A. aberrans Hett. Refrigerated S. Jackson failed
A. croatii Hett. & A. Galloway A. titanum (Becc.) Becc. ex Fresh S. Jackson Failed
Arcangeli
A. dactylifer Hett. A. rostratus Hett. Refrigerated S. Jackson Failed
A. dactylifer Hett. A. rostratus Hett. Frozen S. Jackson Failed
(Continued)

A. dactylifer Hett. A. rostratus Hett. Frozen S. Jackson Failed
A. declinatus Hett. A. gallowayi Hett. Fresh A. Galloway Failed
AGA-2169-01 AGA-2008-02
A. decus-silvae Backer & A. laoticus Hett. Fresh S. Jackson Failed
Alderw.
A. decus-silvae Backer & A. titanum (Becc.) Becc. ex Refrigerated S. Jackson Failed
Alderw. Arcangeli
A. decus-silvae Backer & A. titanum (Becc.) Becc. ex Frozen S. Jackson Failed
Alderw. x A. variabilis Bl.) Arcangeli
A. discophorus Backer & A. laoticus Hett. Fresh S. Jackson Failed
Alderw.
A. dracontioides (Engl.) N. E. Br. (A. decus-silvae Backer & Refrigerated S. Jackson Failed
(Engl.) N. E. Br. Alderw. x A. variabilis Bl.)
A. dracontioides (Engl.) N. E. Br. A. decus-silvae Backer & Refrigerated S. Jackson Failed
(Engl.) N. E. Br. Alderw.
A. dracontioides (Engl.) N. E. Br. A. discophorus Backer & Refrigerated S. Jackson Failed
(Engl.) N. E. Br. Alderw.
A. dunnii Tutch. A. ochroleucus Hett. & V.D. Fresh S. Jackson Failed (3x)
Nguyen
A. dunnii Tutch. A. coaetaneus Liu & Wie Frozen S. Jackson Failed
A. excentricus Hett. A. dactylifer Hett. Frozen S. Jackson Failed
A. galbra F.M. Bailey A. variabilis Bl. ‘Slukas Dark Frozen S. Jackson Failed
Giant’
A. galbra F.M. Bailey (A. variabilis Bl. x A. decus- Fresh S. Jackson Failed
A. gallowayi Hett. A. variabilis Bl. ‘Slukas Dark Refrigerated S. Jackson Failed
Giant’
A. gallowayi Hett. A. barbatus A. Galloway & A. Frozen A. Galloway Failed
AGA-2008-02 Ongsakul AGA-2309-01
A. gallowayi Hett. A. myosuroides Hett. & A. Refrigerated A. Galloway Failed
AGA-2202-01 Galloway AGA-1756-01
A. henryi N. E. Br. (A. variabilis Bl. x A. decus- Fresh S. Jackson Failed
A. henryi N. E. Br. A. bangkokensis Gagn. Fresh S. Jackson Failed
A. henryi N. E. Br. A. laoticus Hett. Refrigerated S. Jackson Failed
A. henryi N. E. Br. A. rostratus Hfett. Frozen S. Jackson Failed
A. hirsutus Teijsm. & Binnend. A. bangkokensis Gagn. Refrigerated S. Jackson Failed
A. hirtus N. E. Br. A. rostratus Hett. Frozen S. Jackson Failed
A. hirtus N. E. Br. AGA-2228-01 A. maxwellii Hett. Frozen A. Galloway Failed
A. hirtus N. E. Br. AGA-2228-01 A. sp. #579 AGA-2176 Frozen A. Galloway Failed
A. impressus Ittenb. A. maxwellii Hett. Refrigerated A. Galloway Failed
AGA-1158-01 AGA-1200-01
A. interruptus Engl. & Gehrm. A. natolii Hett. et al. Frozen S. Jackson Failed
A. interruptus Engl. & Gehrm. A. spec. ‘Pseudodracontium Refrigerated S. Jackson Failed
group’
A. kachinensis Engl. & Gehrm. A. maxwellii Hett. Frozen A. Galloway Failed
AGA-2500-01
A. khammouanensis A. A. konjac K. Koch Frozen A. Galloway Failed
Galloway AGA-2198-01 AGA-2535-01
(Continued)

A. khammouanensis A. kachinensis Engl. & Gehrm. Frozen A. Galloway Failed
A. Galloway AGA-2290-04 AGA-1815-01
A. khammouanensis A. yunnanensis Engl. Fresh A. Galloway Failed
A. Galloway AGA-2290-05 AGA-2506-01
A. konjac K. Koch A. opertus Hett. Refrigerated S. Jackson Failed
A. konjac K. Koch A. titanum (Becc.) Becc. ex Refrigerated S. Jackson Failed
Arcangeli
A. konjac K. Koch A. pygmaeus Hett. Frozen A. Galloway Failed
AGA-1469-01
A. konjac K. Koch A. atrorubens Hett. & Frozen A. Galloway Failed
AGA-1797-01 M. Sizemore AGA-1214-01
A. konjac K. Koch A. gallowayi Hett. Frozen A. Galloway Failed
AGA-1798-01
A. konjac K. Koch A. krausei Engl. Frozen A. Galloway Failed
AGA-1798-01
A. konjac K. Koch A. ochroleucus Hett. & V.D. Fresh A. Galloway Failed
AGA-1798-01 Nguyen AGA-0886-01
A. konjac K. Koch A. longituberosus (Engl.) Frozen A. Galloway Failed
AGA-1947-01 Engl. & Gehrm.
A. krausei Engl. AGA-0283-01 A. hirtus N. E. Br. AGA-2228-01 Fresh A. Galloway Failed
A. krausei Engl. AGA-2619-01 A. maxwellii Hett. Frozen A. Galloway Failed
A. lambii Mayo & Widjaja A. opertus Hett. Refrigerated S. Jackson Failed
A. laoticus Hett. A. bangkokensis Gagn. Fresh S. Jackson Failed (x3)
A. laoticus Hett. (A. variabilis Bl. x A. decus- Fresh S. Jackson Failed
A. laoticus Hett. A. decus-silvae Backer & Refrigerated S. Jackson Failed
Alderw.
A. laoticus Hett. A. haematospadix Hook. f. Refrigerated S. Jackson Failed
A. laoticus Hett. A. henryi N. E. Br. Refrigerated S. Jackson Failed
A. laoticus Hett. A. henryi N. E. Br. Refrigerated S. Jackson Failed
A. laoticus Hett. A. hirtus N. E. Br. Refrigerated S. Jackson Failed
A. laoticus Hett. A. konjac K. Koch Refrigerated S. Jackson Failed
A. laoticus Hett. A. paeoniifolius (Dennst.) Fresh S. Jackson Failed
Nicolson
A. laoticus Hett. (A. lewallei Malaisse & Bamps Fresh S. Jackson Failed
A. laoticus Hett. AGA-1750-01 A. dzui Hett. Frozen A. Galloway Failed
A. laoticus Hett. AGA-2025-01 A. maxwellii Hett. AGA-1200-01 Fresh A. Galloway Failed
A. longituberosus (Engl.) A. variabilis Bl. x A. decus-silvae Fresh S. Jackson Failed (x2)
Engl. & Gehrm. Backer & Alderw.
A. longituberosus (Engl.) A. borneensis (Engl.) Engl. & Frozen S. Jackson Failed
Engl. & Gehrm. Gehrm.
A. longituberosus (Engl.) A. decus-silvae Backer & Refrigerated S. Jackson Failed
Engl. & Gehrm. Alderw.
A. longituberosus (Engl.) A. dracontioides (Engl.) N. E. Br. Frozen S. Jackson Failed
Engl. & Gehrm.
A. longituberosus (Engl.) A. laoticus Hett. Fresh S. Jackson Failed
Engl. & Gehrm.
(Continued)

A. longituberosus (Engl.) A. variabilis Bl. Refrigerated S. Jackson Failed
Engl. & Gehrm.
A. maxwellii Hett. (A. impressus Ittenb. x Refrigerated S. Jackson Failed
A. taurostigma Ittenb. & Hett
‘White Veins’)
A. maxwellii Hett. A. henryi N. E. Br. Fresh S. Jackson Failed
A. maxwellii Hett. A. konjac K. Koch Refrigerated A. Galloway Failed
AGA-1177-01 AGA-2545-01
A. mossambicensis (Schott ex A. maximus (Engl.) N. E. Br. Refrigerated A. Galloway Failed
Garcke) N. E. Br. AGA-0240-01
AGA-0900-01
A. muelleri Bl. A. rostratus Hett. Fresh C. Claudel Failed
A. myosuroides Hett. & A. A. haematospadix Hook. f. Fresh S. Jackson Failed
Galloway
A. myosuroides Hett. & A. A. pygmaeus Hett. Fresh S. Jackson Failed
Galloway
A. myosuroides Hett. & A. A. ongsakulii Hett. & A. Refrigerated A. Galloway Failed (x2)
Galloway AGA-1756-01 Galloway AGA-1534-01
A. myosuroides Hett. & A. (A. variabilis Bl. x A. decus- Frozen A. Galloway Failed
Galloway AGA-1756-01 silvae Backer & Alderw.)
AGA-2642-03
A. myosuroides Hett. & A. A. gallowayi Hett. Refrigerated A. Galloway Failed
A. myosuroides Hett. & A. A. julaihii Ipor, Tawan & P.C. Refrigerated A. Galloway Failed
Galloway AGA-1756-01 Boyce AGA-2811-01
A. myosuroides Hett. & A. A. obscurus Hett. & Refrigerated A. Galloway Failed
Galloway AGA-1756-01 M. Sizemore
A. myosuroides Hett. & A. A. saururus Hett. Fresh A. Galloway Failed
A. natolii Hett. et al. A. sizemoreae Hett. Refrigerated S. Jackson Failed (x3)
A. natolii Hett. et al. A. hirtus N. E. Br. Frozen S. Jackson Failed (x2)
A. natolii Hett.et al. A. titanum (Becc.) Becc. ex Refrigerated S. Jackson Failed (x2)
Arcangeli
A. natolii Hett. et al. A. aberrans Hett. Refrigerated S. Jackson Failed
A. natolii Hett. et al. A. cicatricifer Ridl. Refrigerated S. Jackson Failed
A. natolii Hett. et al. A. hirtus N. E. Br. Refrigerated S. Jackson Failed
A. natolii Hett. et al. A. pygmaeus Hett. Frozen S. Jackson Failed
A. natolii Hett. et al. A. rostratus Hett. Frozen S. Jackson Failed
A. natolii Hett. et al. A. sizemoreae Hett. Frozen S. Jackson Failed
A. natolii Hett. et al. A. tuberculatus Hett. & V.D. Frozen S. Jackson Failed
Nguyen.
A. natolii Hett. et al. A. variabilis Bl. ‘Slukas Dark Refrigerated S. Jackson Failed
Giant’
A. natolii Hett. et al. A. variabilis Bl. ‘Slukas Dark Frozen S. Jackson Failed
Giant’
A. natolii Hett. et al. (A. variabilis Bl. x A. decus- Refrigerated S. Jackson Failed
A. natolii Hett. et al. A. maxwellii Hett. Frozen A. Galloway Failed
AGA-2376-01 AGA-1980-05
(Continued)

A. obscurus Hett. & (A. variabilis Bl. x A. decus- Frozen S. Jackson Failed
M. Sizemore silvae Backer & Alderw.)
A. obscurus Hett. & A. decus-silvae Backer & Refrigerated S. Jackson Failed
M. Sizemore Alderw.
A. obscurus Hett. & A. rostratus Hett. Refrigerated S. Jackson Failed
M. Sizemore
A. obscurus Hett. & A. variabilis Bl. Refrigerated S. Jackson Failed
M. Sizemore
A. obscurus Hett. & M. A. myosuroides Hett. & Fresh A. Galloway Failed
Sizemore AGA-2032-01 A. Galloway
A. ongsakulii Hett. & A. (A. decus-silvae Backer & Refrigerated S. Jackson Failed
Galloway AGA-1534-01 Alderw. x A. variabilis Bl.)
A. ongsakulii Hett. & A. A. gallowayi Hett. Fresh A. Galloway Failed
A. ongsakulii Hett. & A. A. saururus Hett. Frozen A. Galloway Failed
A. ongsakulii Hett. & A. A. titanum (Becc.) Becc. ex Frozen A. Galloway Failed
Galloway AGA-1534-01 Arcangeli
A. ongsakulii Hett. & A. A. obscurus Hett. & Fresh A. Galloway Failed
Galloway AGA-2007-01 M. Sizemore
A. operculatus (ined.) A. interruptus Engl. & Gehrm. Refrigerated S. Jackson Failed
A. operculatus (ined.) A. variabilis Bl. Refrigerated S. Jackson Failed
A. ravenii V. D. Nguyen & A. rostratus Hett. Frozen A. Galloway Failed
Hett. AGA-2179-01 AGA-2166-01
A. reflexus Hett. & A. Galloway A. urceolatus ined. Fresh A. Galloway Failed
AGA-1069-01 AGA-2414-06
A. rostratus Hett. A. laoticus Hett. Refrigerated S. Jackson Failed (x2)
A. rostratus Hett. A. dactylifer Hett. Refrigerated S. Jackson Failed
A. rostratus Hett. A. laoticus Hett. Fresh S. Jackson Failed
A. saururus Hett. AGA-0176-01 A. myosuroides Hett. & Refrigerated A. Galloway Failed (x2)
A. Galloway AGA-1756-01
A. saururus Hett. AGA-0176-01 A. maxwellii Hett. Frozen A. Galloway Failed
A. saururus Hett. AGA-0238-01 A. urceolatus ined. Fresh A. Galloway Failed
AGA-2414-05
A. scutatus Hett. & T.C. A. laoticus Hett. Fresh S. Jackson Failed
Chapman
A. sizemoreae Hett. A. boyceanus Hett. Fresh S. Jackson Failed (x2)
A. sizemoreae Hett. (A. decus-silvae Backer & Refrigerated S. Jackson Failed
A. sizemoreae Hett. (A. gigas Teijsm. & Binnend. x Refrigerated S. Jackson Failed
A. decus-silvae Backer &
Alderw.)
A. sizemoreae Hett. A. gallowayi Hett. Frozen S. Jackson Failed
A. sizemoreae Hett. A. operculatus (ined.) Frozen S. Jackson Failed
A. sizemoreae Hett. A. operculatus (ined.) Refrigerated S. Jackson Failed
A. sizemoreae Hett. A. declinatus Hett. Frozen A. Galloway Failed
AGA-1016-01 AGA-2169-01
A. sp. #587 AGA-2494-02 A. konjac K. Koch Frozen A. Galloway Failed
AGA-0095-01
(Continued)

A. sp. #587 AGA-2494-03 A. ferruginosus A. Galloway Frozen A. Galloway Failed
AGA-2283-01
A. sp. #587 AGA-2494-09 A. hirtus N. E. Br. AGA-2228-01 Fresh A. Galloway Failed
A. taurostigma Ittenb. & Hett (A. variabilis Bl. x A. decus- Refrigerated S. Jackson Failed
‘White Veins’ silvae Backer & Alderw.)
A. thaiensis S.-Y. Hu A. laoticus Hett. Frozen S. Jackson Failed
A. thaiensis S.-Y. Hu A. titanum (Becc.) Becc. ex Refrigerated S. Jackson Failed
Arcangeli
A. thaiensis S.-Y. Hu A. titanum (Becc.) Becc. ex Fresh S. Jackson Failed
Arcangeli
A. thaiensis S.-Y. Hu A. maxwellii Hett. Frozen A. Galloway Failed (x2)
AGA-1928-01
A. thaiensis S.-Y. Hu A. brevipetiolatus A. Galloway, Frozen A. Galloway Failed
AGA-1928-01 A. Ongsakul, & P. Schmidt
AGA-1570-03
A. thaiensis S.-Y. Hu A. titanum (Becc.) Becc. ex Frozen A. Galloway Failed
AGA-1928-01 Arcangeli
A. tuberculatus Hett. & V.D. A. natolii Hett. et al. Frozen S. Jackson Failed
Nguyen.
A. variabilis Bl. A. titanum (Becc.) Becc. ex Frozen S. Jackson Failed (x6)
Arcangeli
A. variabilis Bl. A. cicatricifer Ridl. Refrigerated S. Jackson Failed
A. variabilis Bl. A. laoticus Hett. Frozen S. Jackson Failed
A. variabilis Bl. A. longituberosus (Engl.) Frozen S. Jackson Failed
Engl. & Gehrm.
A. variabilis Bl. A. longituberosus (Engl.) Frozen S. Jackson Failed
Engl. & Gehrm.
A. variabilis Bl. A. natolii Hett. et al. Refrigerated S. Jackson Failed
A. variabilis Bl. A. sumawongii (Bogn.) Bogn.. Refrigerated S. Jackson Failed
A. variabilis Bl. A. thaiensis S.-Y. Hu Frozen S. Jackson Failed
A. variabilis Bl. A. tinekeae Hett. & A. Voge Frozen A. Galloway Failed
AGA-2522-01
A. variabilis Bl. ‘Slukas Dark A. dracontioides (Engl.) N. E. Br. Refrigerated S. Jackson Failed
Giant’
A. variabilis Bl. ‘Slukas Dark A. hirtus N. E. Br. Refrigerated S. Jackson Failed
Giant’
A. variabilis Bl. ‘Slukas Dark A. natolii Hett. et al. Fresh S. Jackson Failed
Giant’
A. variabilis Bl. ‘Slukas Dark A. titanum (Becc.) Becc. ex Refrigerated S. Jackson Failed
Giant’ Arcangeli
A. variabilis Bl. AGA-1245-01 A. crispifolius A. Galloway, Refrigerated A. Galloway Failed
A. Ongsakul, & P. Schmidt
AGA-1753-01
A. variabilis Bl. AGA-1246-01 A. barbatus A. Galloway & Frozen A. Galloway Failed
A. Ongsakul AGA-2309-01
A. variabilis Bl. AGA-1246-01 A. maxwellii Hett. AGA-1200-01 Fresh A. Galloway Failed
A. verticillatus Hett. A. coaetaneus Liu & Wie Fresh S. Jackson Failed
A. verticillatus Hett. A. dracontioides (Engl.) N. E. Br. Fresh S. Jackson Failed
A. verticillatus Hett. A. longituberosus (Engl.) Frozen S. Jackson Failed
Engl. & Gehrm.
(Continued)

A. verticillatus Hett. A. natolii Hett. et al. Refrigerated S. Jackson Failed
A. verticillatus Hett. A. natolii Hett. et al. Refrigerated S. Jackson Failed
A. yuloensis H. Li A. cicatricifer Ridl. Refrigerated S. Jackson Failed
A. yuloensis H. Li A. tuberculatus Hett. & V.D. Fresh C. Claudel Failed
Nguyen.
A. yunnanensis Engl. A. ochroleucus Hett. & V.D. Frozen S. Jackson Failed
Nguyen
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3
Biosynthesis and Decomposition
of Konjac Glucomannan
Orachorn Mekkerdchoo, Yu Lei, and Zhao Jianrong
CONTENTS
3.1 Konjac Glucomannan in the Plant Tissue....................................................................... 102
3.1.1 Characteristics of Konjac Glucomannan Particles............................................. 102
3.1.2 The Formation Process of Konjac Glucomannan............................................... 102
3.2 Biosynthesis of Konjac Glucomannan............................................................................. 102
3.2.1 The Main Carbohydrates in Konjac..................................................................... 102
3.2.2 The Composition of Soluble Glycosides in the Konjac Corm........................... 104
3.2.3 Role of Enzymes...................................................................................................... 105
3.2.4 Biosynthesis Pathway of Konjac Glucomannan................................................. 107
3.3 Enzymatic Degradation of Konjac Glucomannan......................................................... 110
3.4 Regulation of Konjac Glucomannan Synthesis.............................................................. 110
Acknowledgements..................................................................................................................... 112
References...................................................................................................................................... 112
Glucomannan is a cell wall component of many microorganisms and also exists in

plants, such as lily, iris, aloe, mung bean, and pea (Kato et al. 1976; Heller & Villemez
1972; Hinman & Villemez 1975), and also in some species of the Amorphophallus genus
(Liu 2004). The latter are known for being able to synthesize glucomannan, mainly in
the underground tubers. Among them are the main cultivated species, namely, A. konjac
(K. Koch), A. albus (Liu et Chen), and A. muelleri (Bl.). Glucomannan is their main storage
carbohydrate c omponent and accounts for 54% to 61% of the total dry matter. In addition to
glucomannan, the tubers (corms) contain 10%–30% (w/w) starch, 2.6%–7% (w/w) inorganic
elements, 5%–14% (w/w) proteins, 3%–5% (w/w) soluble sugars, and small amounts of
alkaloids (trigonelline) expressed in dry weight (Li et al. 2005).
Since glucomannan is a major component of the tubers of these Amorphophallus species,
it is important to understand its metabolism, as it affects the yield and quality of this crop.
This chapter is to a large extent based on the publication of Liu (2004).
101
3.1 Konjac Glucomannan in the Plant Tissue

3.1.1 Characteristics of Konjac Glucomannan Particles
By cutting off the fresh konjac corms, one can observe many colourless bright granules
with a 0.5 mm diameter that can easily be distinguished from the corm tissue. These gran-
ules are the cells that store the konjac glucomannan. Glucomannan is present in the gluco-
mannan idioblast, which is spherical or elliptical, almost the same size as glucomannan,
and more than 10 times larger than starch granules. In contrast to that, in other plants,
glucomannan is located in the apoplast (the space outside the plasma membrane) area
(Takigami et al. 1997). According to the reports of Ruo Lin Zhong Dao (1957) and Zhao
et al. (2010), the particles of konjac glucomannan stored in the protoplasts (cells without
the cell wall) are flaky, closely adhering to the cell membrane. The viscous glucomannan
particles occupy almost the entire idioblast. In most cases, the particle has a central core
composed of calcium oxalate and liquid cavity mucus. The oxalate crystals are wrapped by
the glucomannan, preventing the tuber from being eaten. The protein membrane of these
cells has to be removed during processing and the needle-shaped oxalate calcium crystals
discarded by sieving (Parry 2009).
3.1.2 The Formation Process of Konjac Glucomannan

When the konjac germination grows, the parenchyma cells of the new corm tissue enter the
differentiation stage, and some of the cells will differentiate into different cells that store
glucomannan. According to a report by Ruo Lin Zhong Dao (1957), in the early stage of
heterologous differentiation, the nucleus is located at the centre of the cell, but it gradually
pushes to the side of the cell due to the increase in the growth of the luminal glycoside, and
oxalic acid is precipitated in the large liquid cavity. Calcium crystallizes and is encapsulated
by liquid cavity mucus, forming the central nucleus of glucomannan particles. Subsequently,
a large amount of glucomannan is synthesized and accumulated around the central core,
and the hetero cells also rapidly expand. Therefore, the growth of the konjac g lucomannan
particles is in a liquid cavity. No increase or decrease in the crystal size of calcium oxalate is
observed during and after the growth of the entire glucomannan particles. When the corm
is germinating again, glucomannan is decomposed for the growth of buds and new corms,
while the central nucleus composed of calcium oxalate or the like inside the glucomannan
particles is discarded with the nutrient-depleted seed (Ruo Lin Zhong Dao 1957).
3.2 Biosynthesis of Konjac Glucomannan

3.2.1 The Main Carbohydrates in Konjac
Tsuji (1895) found that the fine powder extracted from the konjac corm could be detected
by hydrolysis with 3% sulphuric acid, and a large amount of glucomannan was detected.
Therefore, it was first proposed that in addition to starch, the soluble polysaccharide in the
konjac was glucomannan.
Biosynthesis and Decomposition of Konjac Glucomannan 103
Further studies by Mayeda et al. (1911, 1915) confirmed that konjac non-starch polysac-
charide hydrolysate contained glucose in addition to glucomannan, and the ratio of man-
nose to glucose was 2:1.
Goto (1922) named the polysaccharide glucomannan and analyzed the composition of
the carbohydrates in konjac plants. He found that the konjac leaf tissue contained glucose,
fructose, starch, and glucomannan, but no mannose. Glycans accumulated in the leaves,
petioles, and corms, with contents of 19%, 19%, and 40%, respectively.
In view of the above contradictions, Da Yu Hu Nan (1930) performed a detailed experi-
ment and obtained the following results:
1. No free mannose was found in the leaves, petioles, or corms. The presence of glu-
cose, fructose, sucrose, and starch was detected. The content of sucrose and starch
was high. The glucomannan was only found in the corms. This indicated that the
glucomannan in corm is not from free mannan, but it is actually converted from
the accumulated starch in leaves and transferred to the corm to form glucoman-
nan. In addition, there were no significant differences between the results at any
time of the growing season.
2. The starch produced by leaf photosynthesis was converted to sucrose and trans-
ferred to the corms. This can be explained by a higher concentration of sucrose
than other free sugars (glucose and fructose). The concentration of sucrose in dif-
ferent organs is as follows: leaf > petiole > corm. This implies that sucrose can be
transported from the leaves to the corms.
3. The transformed form of glucomannan in the corm was also sucrose. The inter-
conversion of glucomannan and sucrose is a fact that had been established in plant
physiology.
Murata (1972) used Dowex anion exchange resin column chromatography to analyze the
dynamic changes of carbohydrates during the growth of konjac and summarized the
results of the experiments as follows:
1. Sucrose, fructose, and glucose could be detected in any organ, such as seed corm,
growing corms, petioles, and leaf blades. The sucrose content was the highest in
seed corms, growing corms, and leaf blades, and mannose could only be detected
in sprouting seed corms (see Table 3.1).
2. The starch content in the growing corm was maintained at around 0.5% through-
out the growth period (on a fresh weight basis). The content of mannose was lower
in the early stage of growth of the new corm, and gradually increased with the
progressing growth.
Further on, the author systematically analysed the soluble sugars, starch, and glucoman-
nan content of the various organs of konjac (Murata 1975). The results were basically con-
sistent with those reported previously (Murata 1972). With regard to the changes in the
konjac carbohydrates, the following mechanisms are proposed:
1. Leaf photosynthesis products are transported to the corm tissue in the form of
sucrose, and then, in some cells, sucrose is converted to starch; and in other cells,
sucrose is converted to glucomannan.
TABLE 3.1
Changes in Soluble Sugar Content During Konjac Growth (mg/g wet basis)
Growing Total
Organ Days (d) Sucrose Mannose Fructose Glucose Sugars
Seed corm 0 3.61 0.44 0.28 0.24 4.57

20 3.92 0.53 0.33 0.31 5.19
33 5.35 0.87 0.66 0.58 7.46
40 6.64 1.07 2.80 2.40 12.91
47 2.28 0.56 2.25 2.02 7.11
60 0.35 Trace 0.44 0.28 1.07
Growing corm 40 6.40 – 6.35 3.67 16.42
47 3.88 – 3.49 1.62 8.99
60 9.00 – 5.00 4.40 18.40
78 4.70 – 3.95 3.93 17.58
109 2.85 – 3.44 2.06 8.35
139 2.53 – 1.84 0.75 5.16
Petiole 40 1.44 – 4.32 2.80 8.56
47 1.06 – 1.80 1.35 4.21
60 1.97 – 3.32 2.17 7.47
78 1.65 – 3.03 2.12 6.80
109 2.50 – 2.42 1.65 6.57
139 0.80 – 0.10 – 0.90
Leaf 47 3.72 – 1.18 1.34 6.24
60 7.40 – 2.68 1.06 11.14
78 7.80 – 1.14 0.50 9.44
109 12.60 – 1.69 0.34 14.63
139 3.60 – 2.28 0.58 6.46
Source: Murata, T., Nippon Nôgeikagaku Kaishi, 46, 1–7, 1972.
2. Both the starch and the glucomannan can be synthesized in the leaves, but the
content is low, and it is a temporary polysaccharide. The monosaccharides syn-
thesized by photosynthesis in leaves during the daytime can be converted into
polysaccharides, such as starch or glucomannan. Starch and glucomannan are
decomposed into monosaccharides at night-time, while sucrose is synthesized to
facilitate transport.
3. Sucrose, glucose, and fructose are present in both leaves and corms. Glucose and
fructose in the leaves are direct products from photosynthesis, and sucrose is syn-
thesized therefrom. Some of the sucrose in the corm is transported to the leaves,
and then decomposed to form glucose and fructose.
3.2.2 The Composition of Soluble Glycosides in the Konjac Corm

The biosynthesis of glucomannan requires that the monosaccharides involved in the
synthesis be activated first as a nucleoside sugar (or glycoside). Murata (1975) used anion
exchange resin column chromatography and paper chromatography to separate and detect
soluble nucleotides and nucleotide sugars in the acidic ethanol extract of konjac corm,
and identified as Adenosine Diphosphate (ADP) ADP (Adenosine diphosphate)-glucose.

Three kinds of nucleotide sugars were identified, namely, ADP-glucose, GDP (Guanosine
diphosphate)-mannose, and UDP-glucose. Among them, the UDP (Uridine diphosphate)-
glucose content was the highest, while the ADP-glucose content was the lowest. As no
other type of glycosides of mannose was found, it could thus be assumed that the GDP-
mannose was the mannose donor in the konjac glucomannan biosynthesis, similar to
those in microorganisms and mung bean sprouts.
In mung bean glucomannan biosynthesis, the glucose donor is GDP-glucose, but this
glycoside was not found in the konjac corm tissue. Thus, glucose in the konjac glucoman-
nan molecule may be derived from UDP-glucose or ADP-glucose (Heller & Villemez 1972).
Murata (1975) found that the starch synthase isolated from the starch granules of the konjac
tubers was very active when ADP-glucose was used as a glucose donor, while there was no
activity when UDP-glucose was used, i.e., UDP-glucose did not participate or act in the kon-
jac starch synthesis. ADP-glucose was the glucose provider for the konjac starch synthesis.
It was shown that UDP-glucose significantly increased the enzyme activity of the sucrose
synthase crude extract and enhanced the synthesis of sucrose in comparison with ADP-
glucose. UDP was less likely to induce the decomposition reaction of the sucrose synthase
than ADP. Therefore, UDP-glucose and ADP-glucose were both responsible for the sucrose
synthesis reaction and sucrose decomposition reaction of the sucrose synthase, but UDP-
glucose was more involved in the synthesis of sucrose. During the growth of the konjac
tubers, the highly active sucrose synthase contributed to a decomposition reaction, that
was, sucrose was decomposed into monosaccharides, and then used for the synthesis of
glucomannan and starch. The large amount of UDP-glucose inhibited the decomposition
reaction of the sucrose synthase, and, hence, was not conducive to the synthesis and accu-
mulation of polysaccharides. Therefore, it was considered that UDP-glucose did not par-
ticipate in the action of the sucrose synthase in the growing tuber tissue, but as a glucose
donor for the synthesis of glucomannan.
In summary, the photosynthesis products of the leaves are transported to the growing
corms in the form of sucrose. The sucrose is then decomposed into glucose and fructose
by the sucrose synthase and through a series of enzymatic reactions to form activated
monosaccharides: UDP-glucose, GDP-mannose, and ADP-glucose. UDP-glucose and GDP-
mannose are used as glucose and mannose donors, respectively, to participate in the
synthesis of the konjac glucomannan. ADP-glucose acts as a glucose donor for the starch
synthesis.
3.2.3 Role of Enzymes

Konjac glucomannan biosynthesis is a multistep process that involves many enzymes to
convert substrates like sucrose into more complex products, such as glucomannan, accu-
mulated in the konjac corm. The key enzymes in this biosynthesis include the sucrose
synthase, D-mannose-6-phosphate keto-isomerase, and mannose phosphate mutase.
1. Sucrose synthase (Emulsifiable concentrate 2.4.1.13)

The sucrose synthase is the major sucrolytic enzyme. This enzyme converts
sucrose into fructose and UDP-glucose in plants (Geigenberger and Stitt 1993).
Murata (1975) isolated the konjac sucrose synthase for the first time in the study
of soluble nucleotides in the konjac corm tissue. The enzyme has a two-way func-
tion of catalyzing the synthesis and decomposition of sucrose, depending on the
domination by the UDP-glucose or ADP-glucose, respectively. In case the content

of the ADP-glucose is dominating, its main function is to decompose sucrose.
This enzyme is highly active in the growth of the konjac corm tissue.
After the sucrose is decomposed into glucose and fructose, the glucose is further
phosphorylated and activated to produce UDP-glucose or ADP-glucose. Mannose
in the konjac glucomannan is derived from glucose or fructose in the course of the
konjac glucomannan biosynthesis.
2. D-mannose-6-phosphate keto-isomerase (Emulsifiable concentrate 5.3.1.8)
D-mannose-6-phosphate keto-isomerase can be called phosphomannose isom-
erase or mannose isomerase. This enzyme catalyzes the isomerization reaction
between the mannose 6-phosphate and fructose 6-phosphate. The properties
of this enzyme are very similar to those of the enzymes isolated from animals
and yeast. The molecular weight is around 45,000 Daltons, it is stable between
pH 6 and 9, and the enzyme activity is the strongest at pH 6.6 ~ 7.0. The Michaelis
constant (Km) for the mannose 6-phosphate is 0.73 μM, the reaction equilibrium
constant at pH 6.5 is 1.06, and the activation energy of the enzyme is 485.7 kJ/mol.
The enzyme requires divalent metal ions other than Ca2+ and Mg2+, such as Zn2+,
Co2+, Fe2+, Mn2+, or Cu2+ as auxiliary factors. Substances, such as HgCl2 and ethyl-
ene diamine tetraacetic acid, are inhibitors of this enzyme.
Murata (1975) partially purified the phosphomannose isomerase from the kon-
jac corm tissue. The isolation of the phosphomannose isomerase explains the
source of mannose in the konjac glucomannan molecule as being derived from
fructose.
An enzyme capable of converting fructose and mannose to each other is the phos-
phomannose isomerase. It is also present in microorganisms. However, since
Murata (1975) could not detect the presence of this enzyme in the konjac corm
tissue, it is certain that mannose 6-phosphate is not formed by the phosphoryla-
tion of the fructose isoforms to form mannose. Fructose is phosphorylated into
fructose 6-phosphate, which is subsequently converted into mannose 6-phosphate
by the catalysis facilitated by an isomerase.
3. Mannose phosphate mutase (Emulsifiable concentrate 5.4.2.8)
Murata (1976) isolated the mannose phosphate mutase from the konjac corm tis-
sue. The enzyme catalyzes the conversion reaction as follows:
mannose 6-phosphate  mannose 1-phosphate.
The molecular mass of this enzyme is around 62,000 Daltons, it is the most stable
at pH 7.5, and the highest enzyme activity is detected at pH 6.5–7.0. The enzymatic
reaction has an equilibrium constant of 8.5, an activation energy of 46.6 kJ/mol, and
high thermal stability, and Mg2+ and 2-D-glucose-1,6-diphosphate are required as
auxiliary factors. Co2+ or Ni2+ may partially replace Mg2+ for this, while Ca2+ and
Zn2+ are metal inhibitors. Moreover, other enzymes also are found in the konjac
corm tissue (Murata 1976), such as phosphoglucose isomerase that converts man-
nose 6-phosphate and glucose-6-phosphate to each other. Moreover, phosphoglu-
comutase, which catalyzes the conversion of glucose-6-phosphate and glucose
1-phosphate, has also been detected in the konjac corm.
3.2.4 Biosynthesis Pathway of Konjac Glucomannan

The konjac glucomannan biosynthesis pathway was studied for a long time based
on enzyme analysis (Zhang 2004) and molecular analysis by RNA sequencing via 454
DNA deep sequencing with expressed sequence tag analysis (Gille et al. 2011) and de novo
transcriptome with micro Ribonucleic Acid (microRNA) (Diao et al. 2014). These studies
showed that the glucomannan and starch synthesis pathways were correlated, but involved
different types of glucose donors and enzymes responsible for the synthesis.
The photosynthetic products are transported from the leaf to the corm in the form of
sucrose. Under the action of the sucrose synthase, they decompose to form glucose and
fructose. After the action of hexokinase, the glucose and fructose are phosphorylated to
produce glucose-6-phosphate and fructose 6-phosphate. These two phosphorylated mono-
saccharides will be subjected to the catalysis reaction by phosphoglucose isomerase that
interconverts glucose-6-phosphate and fructose 6-phosphate to ensure a balance of glu-
cose and fructose metabolism.
Glucose-6-phosphate is catalysed by phosphoglucomutase to produce glucose
1-phosphate, which is then subjected to ADP-glucose pyrophosphorylase and UDP-glucose
pyrophosphorylase to produce ADP-glucose and UDP-glucose, respectively. As a donor of
glucose in the starch synthesis, ADP-glucose is incorporated into the amylose molecule
by the action of ADP-glucose glycosylase (starch synthase). At the same time, fructose
6-phosphate is catalysed by phosphomannose isomerase to form mannose-6-phosphate,
and then converted to mannose-1-phosphate by phosphoglucomutase, and then GDP-
mannose pyrophosphorylase catalyses the production of GDP-mannose.
Based on enzyme analysis, GDP-mannose acts as a mannose donor, which is involved in
the synthesis of glucomannan under the action of the GDP-mannose transmannosylase.
Meanwhile, UDP-glucose is a donor of glucose, and glucose is added to the glucomannan
molecule under the action of UDP-glucose transglycosylase (see Figure 3.1).
Figure 3.1 shows six enzymes including sucrose synthase, phosphomannose isomerase,
phosphomannose mutase, phosphoglucose isomerase, phosphoglucose mutase, and starch
synthase that have been isolated from the konjac corm tissue. Meanwhile, the other eight
enzymes are currently only assumed and, so far, have not been isolated and identified in
the konjac corms. In addition, the konjac glucomannan molecules are known for branch-
ing, which supposes a branching enzyme and a transacetylase to link some of the glycosyl
groups in the molecule to the branches and acetyl groups. Yet, there does not appear to be
published reports about the isolation of these enzymes from the konjac corm.
For molecular analysis, Gille et al. (2011) constructed a complementary Deoxyribonucleic
Acid (cDNA) library from 454 RNA sequencing using the developmental stage of the A. kon-
jac corm, and then blasted with known gene encoding proteins in the sugar conversion path-
way of Arabidopsis, which can produce glucomannan in the cell wall. This method allowed
them to identify the important role of an enzyme in the glucomannan synthesis pathway
based on transcription (high encoding proteins abundance). The glucomannan biosynthe-
sis pathway could be re-constructed as shown in Figure 3.2. The sucrose synthase, UDP-D-
glucose pyrophosphorylase, phosphoglucomutase, and all enzymes in starch synthesis are
major pathways with high encoding proteins reads, while hexokinase and phosphofructoki-
nase are minor pathways. This indicates that fructose had a minor role in this pathway.
Glucomannan can be synthesized from GDP-D-mannose and GDP-D-glucose by
the cellulose synthase-like family A (CSLA) as glucomannan synthase. Proteins of the
family CSLA belong to the glycosyltransferase family 2. In Arabidopsis, CSLA3 is involved
in glucomannan biosynthesis in secondary cell walls (Goubet et al. 2009). However, the
FIGURE 3.1
Schematic diagram of glucomannan and starch biosynthesis based on enzyme analysis. AGP: ADP-glucose
pyrophosphorylase; CSLA: cellulose synthase-like A; CSLD: cellulose synthase-like D; GMPP: GDP-mannose
pyrophosphorylase; HXK: hexokinase; INV: invertase; PGI: phosphoglucose isomerase; PGM: phosphogluco-
mutase; PMI: phosphomannose isomerase; PMM: phosphomannose mutase; SBE: starch branching enzyme; SS:
starch synthase; SuS: Sucrose synthase; UGP: UDP-glucose pyrophosphorylase. (Modified from Zhang, X.G.,
Biosynthesis of konjac glucomannan, in: Konjac Biology, ed. P. Y. Liu, pp. 84–91, China Agriculture Press, Beijing,
China, 2004; Diao, Y. et al., PloS One, 9, e95428, 2014.)
synthesis pathway of GDP-D-glucose, which is accepted by CSLA glucomannan syn-

thases with GDP-D-mannose (Liepman et al. 2005), is still questionable, as the GDP-D-
pyrophosphorylase for the synthesis of GDP-D-glucose from glucose-1-phosphate has
not been described in the konjac so far. The alternative pathway would be converting GDP-
D-mannose into GDP-D-glucose by GDP-D-mannose-2-epimerase (Reiter 2008). Moreover,
the CSLA3 protein was recombined in yeast Pichia pastoris for activity confirmation.
The result showed that Pichia cells were significantly expressing the CSLA3 protein by
more GDP-glucose and GDP-mannose production than wild-type samples.
Diao et al. (2014) studied microRNA and transcriptome analysis in the leaves of A. konjac
and A. bulbifer. This study revealed the CSLD for the first time, indicating the corresponding
genes in both konjac species. CSLD proteins would be the glucomannan synthases that are
using UDP-glucose and GDP-mannose as substrates (blue line in Figure 3.2). Moreover,
A. bulbifer had a lower expression level in ADP-glucose pyrophosphorylase, but a higher
level in UDP-glucose pyrophosphorylase than A. konjac. This reflects the high amounts
of glucose-1-phosphate which correspond to glucomannan synthesis, as A. bulbifer had a
higher glucomannan content than A. konjac.
Biosynthesis and Decomposition of Konjac Glucomannan
FIGURE 3.2
Suggested glucomannan and starch synthesis pathway based on EST database in A. konjac corm. (Modified from Gille, S. et al., Planta, 234, 515–526, 2011; Diao, Y. et al.,
PloS One, 9, e95428, 2014.)
109
There are various studies claiming that CSLA proteins exhibit mannan synthase a ctivity.
The CSLA protein can transfer mannosyl residues from GDP-mannose to produce a homo-
mannan backbone (Sawake et al. 2015). CSLA2, CSLA3, and CSLA9 are responsible for the
synthesis of the glucomannan in Arabidopsis stems, and CSLA7 synthesizes the glucoman-
nan in embryos, according to Goubet et al. (2009). CSLA2 is also involved in the biosyn-
thesis of the mucilage glucomannan in the structure of the Arabidopsis seed (Yu et al. 2014).
Arumingtyas and Fatinah (2015) designed a primer from the CSLA gene to detect its influ-
ence on the glucomannan content of A. muelleri from Java Island, Indonesia. However, this
relationship could not be found.
3.3 Enzymatic Degradation of Konjac Glucomannan

After the konjac sprouts, the glucomannan, which is the main storage material, will be
gradually degraded by enzymatic action, and the degradation products will be trans-
ported to the new buds and corm tissues to serve as a carbon source and energy source for
the enhancement of new cells.
Mayeda (1920) and Ohtsuki (1930) pioneered the use of the enzymes in snails and alfalfa
to hydrolyse glucomannan. The mannanase, which separates the hydrolyzable mannan
from microorganisms, has also been used for the enzymatic studies of the konjac gluco-
mannan. Until the 1970s, Sugiyama et al. (1973) isolated and purified the mannanase from
the germinated corm tissue. The enzyme preparation was very stable between pH 4.0 and
8.0. The optimum enzyme activity was at pH 4.7, and the optimum reaction temperature
was 40°C. Above 50°C, the enzyme activity decreased rapidly. The enzyme degrades the
konjac glucomannan following a random pattern. In the digestive products of the mannan
by the mannanase, three monosaccharides and at least six oligosaccharides were detected.
Of the three monosaccharides, one was glucose and the other was mannose. The oligo-
saccharides are further broken down into monosaccharides, and several enzymes are
required to participate. Therefore, the decomposition of the konjac glucomannan is a com-
plex process involving a variety of enzymes mainly composed of mannanase.
3.4 Regulation of Konjac Glucomannan Synthesis

The konjac glucomannan content is an important indicator of the konjac quality. One of the
desirable characteristics of the ideal konjac variety is little starch, but mostly glucomannan.
Using the conventional breeding techniques, it is difficult to significantly increase the con-
tent of the konjac glucomannan. In the konjac flour obtained from the mature corms, the
starch content accounts for about 10% to 20% of the dry weight and is removed during
processing. By incorporating the knowledge of mechanisms of the biosynthesis of the kon-
jac glucomannan in the breeding process in order to reduce the synthesis of starch in the
konjac tuber tissue, the glucomannan content of the tubers could be increased. As shown
in Figure 3.1, the ADP-glucose pyrophosphorylase and starch synthase are closely related
to the starch synthesis. If antisense gene inhibition technology is used, the synthesis of
starch could be blocked.
ADP-glucose pyrophosphorylase is a key enzyme in the starch biosynthesis. The enzyme

is a heterotetramer composed of two subunits of different size. The small subunit is
extremely conservative in evolution and is encoded by the same gene in the photosyn-
thesis and non-photosynthetic tissues. It is currently believed that small subunits play a
major role in the catalytic reaction. Therefore, separating the small subunit of the enzyme
and then introducing it into the konjac genome would inhibit the expression of the small
subunit gene.
In order to achieve this objective, the following procedure would have to be applied:
First, extract RNA from the growing konjac corm tissue, and synthesize the single-
stranded cDNA using reverse transcriptase. Then apply the primers and Polymerase
Chain Reaction (PCR) designed for the functionally conserved segment of the ADP-glucose
pyrophosphorylase small subunit that is isolated from other organisms. In this technique, a
part of the cDNA fragment is amplified in a large amount and inserted into the downstream
of the promoter of the plant expression vector in the reverse manner of the transcription
of the original gene to construct an antisense expression vector. After the antisense gene
has been introduced into the konjac, an mRNA complementary to the normal ADP-glucose
pyrophosphorylase small subunit gene transcription product is transcribed, and the two
mRNAs hybridize by complementation, thereby preventing a ADP-glucose pyrophospho-
rylase small subunit. The base messenger Ribonucleic Acid (mRNA) is translated into a
protein to achieve the purpose of reducing the production of the enzyme and inhibiting the
activity of the enzyme. The inhibition process of the antisense gene is shown in Figure 3.3.
FIGURE 3.3
Antisense gene expression inhibition process. (Modified from Zhang, X.G., Biosynthesis of konjac g
lucomannan,
in: Konjac Biology, ed. P. Y. Liu, pp. 84–91, China Agriculture Press, Beijing, China, 2004.)
Alternatively, controlling the sucrose degradation and sucrose synthase production at a

transcription level are also an option. For example, microRNA 339 can induce the silencing
of the sucrose synthase mRNA by combining with the transcripts of the sucrose synthase
(Ohtsuki 1930). In the leaves, fructose and glucose can be obtained by sucrose decomposi-
tion and also can be directly produced by photosynthesis, providing the raw materials
of the glucomannan and starch synthesis. However, fructose and glucose in the corms of
Amorphophallus are mainly obtained from sucrose decomposition. Therefore, the suppres-
sion of the sucrose synthase in the leaves possibly enhances the use of photosynthesis-
produced glucose and fructose in the starch and glucomannan synthesis.
Acknowledgements
The authors would like to acknowledge the assistance of Dr. Pilanee Vaithanomsat,
Enzyme Technology and Waste Management Research Unit, Kasetsart Agricultural and
Agro-Industrial Product Improvement Institute (KAPI), Kasetsart University Bangkok, for
the critical review of this chapter.
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4
Field Production of Konjac
Zhang Shenglin, Jiang Xuekuan, and Hadi K. Purwadaria
CONTENTS
4.1 Cultivation........................................................................................................................... 116
4.1.1 Variety Improvement of Konjac............................................................................ 116
4.1.1.1 Current Konjac Varieties Cannot Meet Production Needs................ 116
4.1.1.2 Objectives of Variety Improvement....................................................... 117
4.1.1.3 Basis of Variety Improvement................................................................ 117
4.1.2 Cultivated Species and Varieties.......................................................................... 120
4.1.2.1 Cultivated Species and Varieties in China........................................... 121
4.1.2.2 Cultivated Species and Varieties in Japan............................................ 125
4.1.2.3 Cultivated Species and Varieties in Other Countries and Regions...... 129
4.1.3 Elite Breeding of Konjac......................................................................................... 130
4.1.3.1 Establishment of Elite Variety Breeding Grounds for Konjac........... 130
4.1.3.2 Principles of Elite Breeding System for Konjac.................................... 131
4.1.3.3 Use of Sterilized Konjac Seed Material................................................. 132
4.1.3.4 Local Production and Commercialization of Elite Varieties............. 132
4.1.4 Ecological Adaptability of Konjac........................................................................ 133
4.1.4.1 Influence of Temperature on Growth and Development of Konjac..... 133
4.1.4.2 Influence of Light on Growth and Development of Konjac............... 134
4.1.4.3 Influence of Moisture Conditions on Growth and Development
of Konjac.................................................................................................... 135
4.1.4.4 Mineral Nutrition..................................................................................... 136
4.1.4.5 Soil.............................................................................................................. 137
4.1.4.6 Other Conditions...................................................................................... 137
4.1.5 Hazards for Konjac Cultivation............................................................................ 138
4.1.5.1 Diseases..................................................................................................... 138
4.1.5.2 Insect Attacks on Konjac......................................................................... 140
4.1.5.3 Meteorological Hazards.......................................................................... 140
4.1.5.4 Drought Damage...................................................................................... 141
4.1.5.5 Waterlogging Damage............................................................................. 141
4.1.5.6 Damages Caused by Wind and Hail..................................................... 141
4.1.6 Cultivation Techniques.......................................................................................... 142
4.1.6.1 Selection of Areas and Plots for Cultivation........................................ 142
4.1.6.2 Cultivation Pattern and Cropping System........................................... 144
4.1.6.3 Reproduction Methods............................................................................ 146
4.1.6.4 Seed Konjac Selection and Treatment Before Planting....................... 147
4.1.6.5 Plot Preparations...................................................................................... 148
115
4.1.6.6 Planting..................................................................................................... 148

4.1.6.7 Field Management................................................................................... 149
4.1.6.8 Harvesting................................................................................................ 150
4.1.6.9 Storage of Planting Material................................................................... 150
4.2 Agroforestry-Based Production System in Indonesia................................................... 152
4.2.1 The Agroforestry Environment and Its Sustainability..................................... 152
4.2.1.1 Konjac Agroforestry System and Environment................................... 152
4.2.1.2 Agroforestry Sustainability.................................................................... 153
4.2.2 Konjac Growth Cycle.............................................................................................. 154
4.2.2.1 Cultivation................................................................................................. 154
4.2.2.2 Harvesting of Tubers............................................................................... 154
4.2.2.3 Replanting................................................................................................. 155
4.2.3 Socio-Economic Aspects of Agroforestry............................................................ 156
References...................................................................................................................................... 156
4.1 Cultivation
4.1.1 Variety Improvement of Konjac
According to the evolutionary process of various Amorphophallus species, the konjac is a
plant with a long history. Although the konjac has been cultivated and consumed for a
very long time in a number of regions in China, Japan, and Southeast Asia, the plant is still
grown under semi-wild conditions in many producing areas. The crop is cultivated from
unimproved original genetic material in most regions and cannot meet the demand for
industrial-scale production.
4.1.1.1 Current Konjac Varieties Cannot Meet Production Needs

The dormancy period of the konjac is as long as half a year. Although it is planted in soil
for half a year, during April–June, the konjac is in a budding state, coming up out of the
ground, undergoing leaf expansion, and the ‘head-changing’ stage in order to get prepared
for yield formation. During July–September, the konjac really enters the stage of the inten-
sive production of nutrients and accumulation of reserve substances. The photosynthetic
efficiency of the konjac is lower than many other tuber crops, and the optimum tempera-
ture is also low, so is the efficiency of producing and accumulating of the reserve substances
(Liu Peiying 2004). Besides, the konjac has strict requirements for ecological conditions.
The outbreak of bacterial soft rot (especially Erwinia cartovorum) and fungal southern blight
(Sclerotium rolfsii) may be caused if the environmental conditions for cultivation are unsuit-
able (very wet soil), causing a severe drop in the yield or even a total crop failure.
Potatoes can grow more than ten new tubers and can be cut into pieces for reproduction.
However, a single mother corm can grow only one daughter corm with 5–8 buds. Hence,
the konjac cannot be cut into pieces for reproduction like potatoes; and this biological fea-
ture is stable and cannot easily be genetically changed. Therefore, the reproduction coef-
ficient of the konjac is very low.
In conclusion, both the yield formation ratio and reproduction ratio of the konjac are
lower than in most other tuber crops. This is why most varieties of the konjac cannot meet
the production needs.
Field Production of Konjac 117
4.1.1.2 Objectives of Variety Improvement
1. To breed varieties with strong resistance to soft rot and southern blight.
2. To breed varieties with corms characterized by high dry matter weight.
3. To breed varieties with relatively shorter dormancy period and longer period from
leaf expansion to the maturity stage.
4. To breed varieties with strong adaptability to light and temperature and high pho-
tosynthetic efficiency.
5. To breed varieties with the appropriate ratio of the number of buds to corm.
For species with few buds, such as the Amorphophallus corrugatus N. E. Brown (syn.
A. tianyangense P. Y. Liu & S. L. Zhang), the varieties with more buds should be
bred, and for species like the A. albus with too many buds, the varieties with more
developed corm and less buds should be bred.
6. For species producing glucomannan, the varieties with high glucomannan content
(KGM), high viscosity, and white colour should be bred. For species producing
mainly starch, the varieties with a high starch content and suitability for particu-
lar applications should be bred.
4.1.1.3 Basis of Variety Improvement

4.1.1.3.1 Collection and Collation of Genetic Resources
According to the latest statistics of the International Aroid Society, there are a total of
167 species of the Amorphophallus genus. In recent years, the collection, collation, and
study of the genetic resources of the Amorphophallus have been carried out at Southwest
University in Chongqing, Yunnan Academy of Agricultural Sciences, Enshi Prefecture
Academy of Agricultural Sciences, Hubei Province, and other institutes in China.
Researchers in Japan, Australia, and other countries have also gradually conducted
investigations and studies of the genetic resources of the konjac. There are 21 species
of the Amorphophallus that are growing in China. However, so far, only their botanical
characteristics are known. The systematic research and development of konjac have been
conducted on A. konjac and A. albus; however, the comprehensive collection and study
of the economic characters, adaptability, and disease resistance have not been carried
out on other species (Li Heng 1979). Studies on the A. muelleri (including A. bulbifer and
A. yuloensis), A. ximengensis (syn. Amorphophallus krausei), A. paeoniifolius, and other spe-
cies have been carried out in recent years. The konjac resources in Japan are relatively
limited and mainly include the A. konjac. In addition to the native Japanese varieties,
Chinese A. konjac resources have also been introduced. The crossbreeding of the A. konjac
has been carried out. The konjac in Vietnam, Burma, Thailand, Laos, the Philippines,
Indonesia, and other Southeast Asian countries is in a transient stage from wild genetic
resources to domestication. Although the konjac is cultivated in India, the main species
is the starch type A. paeoniifolius. Due to factors such as the wide range of centres of
origin, more countries in which the konjac is grown, and more specific climatic require-
ments of the konjac than those of other crops, there is still insufficient knowledge about
the genetic resources of various species of the Amorphophallus. Before collection, colla-
tion, and systematic studies of representative specimens have been carried out, we are
still far away from meeting the demand for resource development, domestication, breed-
ing, and field cultivation.
4.1.1.3.2 Basis for Innovative Development of Genetic Resources

The basic research includes the introduction of varieties to new locations, identification of
genetic relationships, propagation of particular species, and study of interspecific incom-
patibility. Natural genetic resources are shared by mankind. Therefore, it is necessary to
carry out the study of the genetic resources of the konjac on a collaborative level through-
out the world. The Japan Gunma Prefecture Comprehensive Agriculture Proving Ground
has carried out a series of variety improvement projects by introducing the Chinese A.
konjac germplasm and crossbreeding with the native Japanese A. konjac germplasm
(GPAIA 1991). On the basis of the collection and introduction of the domestic konjac germ-
plasm, Southwest University in Chongqing, Enshi Prefecture Academy of Agricultural
Sciences in Hubei Province, Yunnan Academy of Agricultural Sciences, and other insti-
tutes in China also introduced the konjac germplasm from Japan, Thailand, Burma,
and other countries and carried out subsequent variety improvement. Vietnam, Burma,
Thailand, Laos, the Philippines, Indonesia, and other Southeast Asian countries are the
centre of origin and have rich genetic resources of the konjac, but are still at the early stage
of the systematic collection, collation, and study of such resources.
Researchers have conducted karyotypes analysis of such genetic resources in China as
the A. konjac, A. albus, A. kiusianus, A. pingbianensis, A. yunnanensis, A. yuloensis, and A. kachi-
nensis and have preliminarily determined the Giemsa-c banding pattern of the above spe-
cies (Liu Peiying et al. 1985). On the basis of gel electrophoresis, the genetic relationship
among genetic resources was conducted. The results indicated that the genetic relation-
ship among the germplasm of the konjac is basically consistent with its morphology and
geographical distribution and can be used as an important reference for crossbreeding
(Zhang Yujin et al. 2001). Studies have shown that the traits of the F1 generation of the kon-
jac are related to the maternal parent, and that such generation has strong heterosis, can
often concentrate the good traits of its parents, and can be reproduced asexually. An exam-
ple is the Nonglin 2 (Akagi-ohdama) bred in Japan. Its female parent is a ‘Chinese variety’
(Shina), and its male parent is a Japanese ‘native variety’. Its F1 generation that reproduced
asexually has stable traits: the part above the ground is like its female parent, but its lateral
buds look like its male parent, and it has a long spherical shape, however, compared with
its male parent, the surface grooves are fewer, the epidermis colour is slightly lighter, and
the underground part is close to an intermediate type between its parents. Other research
reports have shown that the traits of the F1 generation also resemble those of the female
parent, for example, A. albus (♀) × A. konjac (♂). The plant of the F1 generation is small; and
the colour of petioles and the plant architecture are like those of the A. albus. For A. konjac
(♀) × A. albus (♂), the plant of their F1 generation is tall; and the colour of petioles and the
plant architecture are like those of the A. konjac (Zhang Shenglin et al. 1998). The core par-
ent materials should be selected, identified, and cultivated in order to maintain a wide
range of genetic resources according to the breeding objective. As the only cultivated spe-
cies in Japan, the A. konjac shows a good adaptability and can be used as the best and the
most important core parent material for cultivation. The A. albus is a species known for a
high Konjac Glucomannan (KGM) content and can be well used as the core parental mate-
rial for cultivation. Although the yellow konjac (various species, see section 4.1.2) has a
lower content of KGM than the aforementioned two species, it is highly heat resistant and
can be used in breeding. The A. paeoniifolius and A. dunnii can be used as the core parents
for the cultivation of starch type konjac varieties; and their genes for disease resistance and
stress resistance need to be thoroughly researched and developed in combination with
modern biotechnology.
4.1.1.3.3 Review of Breeding Methods

4.1.1.3.3.1 Plant Selection and Cultivation of Intraspecific Varieties Since some Amorphophallus
species are prone to hybridization, a rich intraspecific variation appeared in the long pro-
cess of natural evolution. Such variation may be reflected in the significant differences in
the characteristics of a single plant, or may be stable and consistent to differentiate groups
of plants, providing a basis for plant selection. Plant selection is essential for the breeding
of varieties resistant to diseases and stress. For instance, the A. albus has such types as
the ‘green type’ with pure green petioles, the ‘spotted type’ with petioles with brownish
green spots, and the ‘yellow type’ with petioles with a ground colour of yellow in different
regions or in the same region, and has excellent yielding ability, disease resistance, and
adaptability and other desirable attributes (Sun Yuanhang 2006). The intraspecific farm-
grown species of the A. konjac in different regions also show variation in different char-
acteristics. For example, the buds of some A. konjac groups detach from the mother corm
and shrivel into a garlic head-like shape; the plants of some A. konjac lines are compact and
have a low degree of expansion; and some A. konjac lines have many split leaves with a
high degree of expansion. On this basis, there are significant differences in petiole colour
and spots of the A. konjac in different regions, which are the most noticeable traits for vari-
ety breeding.
4.1.1.3.3.2 Mutant Breeding The asexually reproduced crops show differences in charac-
teristics caused by mutations. A suitable variation is obtained by artificial selection and
multiplied by asexual reproduction. This has been an important way of tuber crops breed-
ing. The physical and chemical methods are the most common means of mutant breeding.
In mutation breeding by irradiation, the selection of the appropriate irradiation dose is the
condition of successful mutation. If the dose is too small, it is insufficient to cause variation
or the variation is not significant. If the dose is too high, the variant cells may have reduced
vitality and even die; and even if such cells survive, inferior mutations will be caused and
cover up other mutations. When the konjac corm is planted after the irradiation treatment,
it gradually shrivels and disappears at the early stage of the growth. The new corm and
its buds can be regarded as the M2 generation. Whether the superior mutations are con-
solidated by the M3 generation can be observed and recorded. A new variety or strain can
be bred by asexual reproduction. The meristem of the konjac is at the end where the ter-
minal bud is located, i.e., the upper end of the corm. The lower end, however, is the stor-
age tissue. The mutation treatment usually must be centred on the terminal bud of the
corm. After the mutation treatment, the plants and new corms grown in the same year
can be observed to see if the variation has occurred and first assess whether it is a favour-
able variation. However, since the konjac corm is very crisp and tender and the radiation
may cause injury by infection through diseases or become fruitless after the injury, large
corms cannot be used as the material for the mutation treatment; instead, corms with low
water content and a single corm weight of no more than 50 g should be used. Therefore, the
corms grown in the same year are insufficient to represent the content of the KGM. Such
corms should be cultivated for another year until they reach the size of the commercially
grown konjac. The effects of irradiation on the content of the KGM can be explained cor-
rectly by the analysis conducted at this time (Huang Xunduan 2002). The common chemi-
cal mutagens include gibberellin and colchicine. It has been reported that on the basis of
the liquid culture of the konjac, polyploids are obtained by inducing the meristem of the
A. konjac with colchicine, in order to conduct the test on the polyploid breed of the konjac
(Hu Nan 2011).
4.1.1.3.3.3 Crossbreeding The difficulties in the crossbreeding of the konjac are mainly
reflected in the following aspects:
1. Few materials to choose from: only a few varieties, such as the A. albus, A. konjac,
A. muelleri, A. yunnanensis, and A. krausei, can be used as the materials for cross-
breeding, and these varieties account for a small proportion of the genetic
resources of the konjac in the world. Therefore, the genetic background is narrow
for breeding materials.
2. Difficulties in hybridization technique: there are such problems as difficult flower
induction, different flowering periods, and self-incompatibility. It takes 3–4 years
from seed to blossom of the konjac, the flowering ages of different varieties are
different, completely different flowering periods of the same konjac plant can be
caused due to the difference in environmental conditions, in addition, the changes
in the temperature and humidity in the environment not only affect the konjac
blooms, but also have great impacts on anther dehiscence, no pollen spread, and
stigma activity. The konjac is a plant with the same inflorescence of the pistil and
stamen and different flowers. The pistil matures 1–2 days earlier than the stamen
in the same inflorescence, and the suitable time of the pistil for fertilization is gen-
erally as short as 1 day. If the odour is released from the appendage, it indicates
that the pistil is mature. The effective pollination period of the konjac is from the
time of the pollen spread to the maturity of the stamen and is about 1–2 days.
When the pollen is spread by the stamen, the pistil stigma of the same plant has
lost its pollination and fertilization vitality. Therefore, a certain group is needed
for cross-pollination to ensure that the pistil and stamen mature at the same time
during the intraspecific hybridization of the konjac. The flowering periods of dif-
ferent varieties of the konjac are very different and may be at any time during
March–September, which is also a major obstacle to the interspecific crossbreeding
of the konjac. The manipulation of the temperature and humidity and the induc-
tion by exogenous hormones has provided preliminary solutions to the problems
in the crossbreeding of the konjac, such as induced flowering, control of flowering
period, cross-incompatibility, and pollen preservation (Zhang Shenglin 1995).
4.1.1.3.3.4 Breeding Using Biotechnological Techniques In the last 20 years, the application of
biotechnology in a variety of improvements has made significant progress and has been
applied in field production. However, there are few reports on the applications of biotech-
nology in breeding of the konjac. The establishment and perfection of tissue culture for
the konjac laid the foundation of biotechnological breeding for overcoming the incom-
patibility of sexual hybridization by cell fusion. Zhang Xingguo et al. (1992) explored the
cell fusion of the konjac and completed the process from the protoplast isolation of the
petiole cells to the somatic cell hybridization of the embryogenic cell and fusion cell; and
the plant regeneration rate is up to 62%. The genetic engineering breeding of the konjac is
still at a basic research stage. The current reports are mainly on the research of a disease-
resistant gene, heat-resistant gene, high-yielding gene, KGM synthesis, storage gene, and
anti-browning gene (Bai 2016).
4.1.2 Cultivated Species and Varieties

The Amorphophallus species are divided into starch or KGM types according to their main
component. The main component of the starch type species is starch almost without the
KGM. Meanwhile, since the corm contains a toxic alkaloid, the starch type konjac had
not been valued and developed by mankind in the long history of frequent food short-
ages. Although the KGM type konjac has high-quality soluble dietary fibres, traditional
processing just makes it into konjac gel food by adding alkali to satisfy hunger in famine
years or act as an off-season specialty. From the perspective of agronomy, the konjac is still
a very basic crop so far; Japan was the first country to carry out the selection and breeding
of the konjac; and China has so far successively bred several varieties in recent years, but
many producing areas in Southeast Asia have not yet reached the cultivation stage and
still harvest it mainly in the wild.
4.1.2.1 Cultivated Species and Varieties in China

China was the first country to cultivate and consume the konjac. Although its cultivation
and utilization has lasted for more than a thousand years, this crop was recorded in his-
tory mostly in times of food shortages to satisfy hunger, and its value was not recognized
by people. Therefore, its nutritional properties, seed selection, and breeding of suitable
varieties had not attracted people’s attention. It was only during the recent 40 years that
China has gradually implemented breeding programs of new varieties.
China is the host of 21 konjac species, most of which are of the KGM type. The A. konjac
accounts for more than 90%, followed by the A. albus. Each variety has a certain distribu-
tion range and presents many variations after a long time of natural and artificial selec-
tion. In the last 20 years, purposeful selection and breeding, hybridization, and mutation
have been carried out. The main cultivated species and selected varieties are as follows.
4.1.2.1.1 A. albus
The A. albus (P. Y. Liu & J. F. Chen) was discovered and named by Liu Peiying and Chen
Jinfeng from Southwest University in 1984. This species is naturally distributed in the
Jinsha River valley, and the main area suitable for this variety is the downstream sec-
tion of the river valley at the border between southern Sichuan Province and northern
Yunnan Province. This species produces a small plant, with a green three-lobed leaf, that
grows up obliquely and has fewer split leaves than the A. konjac. The corm is oblate with
a sunken neck, brown epidermis, white interior, and developed offset tubers. There is a
sterile neuter between inflorescences. According to the system identification, the A. albus
is an endemic species of China. Its content of KGM can be up to 60%. The molecular weight
of the KGM is also greater than that of the KGM in the corm of the A. konjac. The A. albus
is a KGM type konjac with the best processing quality (Liu & Chen 1984). The browning
during the processing of the A. albus is slight. The konjac flour produced from its corm is
white in colour and has a much better viscosity, transparency, and other quality attributes
than the flour from the A. konjac.
The A. albus can adapt to an environment with low altitude, high temperature, low
humidity, and strong sunlight. Its active accumulated temperature is 4863.1°C and effec-
tive accumulated temperature is 1658.1°C. After years of natural selection and domestica-
tion, the suitable stable variations occurred in different cultivation areas, forming local
germplasm with different colours and spots and also significantly different yield and
content of the KGM. The ‘green type’ with pure green petioles, ‘spotted type’ with peti-
oles with brownish green spots, and ‘yellow type’ with petioles with a ground colour of
yellowish-brown are superior strains of the A. albus. The A. albus has a good quality of
the KGM and strong resistance to soft rot and southern blight. However, the single plant/
unit area yield is low; the offset tubers account for a too large proportion of the weight
of the underground harvested part; and can be more than 50% in particularly fertile soil.
Therefore, the blind introduction of the A. albus is not advocated. Meanwhile, seed mate-
rial selection and breeding should be rigorously carried out on the A. albus in order to
maintain its merits, such as good resistance to diseases, high reproduction coefficient, and
high quality, in order to offset its demerit of a low yield.
4.1.2.1.2 A. konjac
A. konjac (K. Koch) is distributed in all producing areas in China and is the most impor-
tant cultivated species. The A. konjac has good adaptability, a wide range of suitable areas,
and can adapt to all mountainous areas suitable for the konjac in China. The A. konjac
can also grow well in areas specialized in producing the A. albus, especially in areas
with a northern latitude, high altitude, low temperature, weak sunlight, and high humid-
ity. From germination to senescence, the A. konjac needs an active accumulated tempera-
ture of 4278°C (above 10°C) and an effective accumulated temperature of 1089.3°C (above
15°C). Characteristics of the A. konjac are: nearly circular corm, reddish-brown epider-
mis, black brown small spots, and large size; bright pink terminal bud of the corm and
inverted ‘V’-shaped leaf bud with a length of only about 1 cm; and an inverted ‘U’-shaped
flower bud with a length of more than 1.5 cm and up to 10 cm before spring planting.
The flower bud can be easily recognized for early treatment. The A. konjac has obvious
apical dominance. The offset tubers are concentrated on the middle-upper part of the
corm. The 2-year-old plants have strong growth vigour and a plant height of 1–1.5 m.
The ground colour of the petiole is related to age. The morphologic characteristics of 2- to
3-year-old plants are stable; the petiole is green or light green in the ground colour; and
it has connected olive-drab spots with small white dots, covering the petiole. The spathe
is funnel-shaped, with a coiled base. The surface of the spathe is pale green and has dark
green spots. The interior is deep purple-red. The inflorescence is one time longer than
the spathe. The pistillate inflorescence is cylinder-shaped; and the appendage is sword-
shaped, purple-red, and hollow. The filaments are united with a four-chamber anther
and spherical pollen; the stigma is trifid or bifid with papillary protuberance. The fruit
chamber is oval and first is green and then turns orange when it matures. The flowering
period is April–May, and the fruiting period is June–August. The A. konjac can adapt to
the environmental conditions of high altitude, low temperature, and insufficient sunlight.
In recent years, the in-depth innovation of genetic resources and variety improvement of
the A. konjac have been carried out in various regions. Several varieties of the A. konjac
have been successively bred:
4.1.2.1.2.1 ‘Wanyuan A. konjac’ In the late 1980s, on the basis of the collection of the
genetic resources of konjac in China, the former Southwest Agricultural University (now
Southwest University) in Chongqing optimized 15 strains of the cultivated specimens of
the A. konjac from various regions. After systematic breeding and 4 years of variety com-
parison tests and regional tests, a new variety was approved by the Sichuan Crop Varieties
Examination and Approval Committee as ‘Wanyuan A. konjac’ in 1993. The variety is
characterized by a strong growth vigour, good resistance to stress, and high yield. It was
not only the first approved performing variety of the A. konjac in China, but also became
the dominant variety in the Daba Mountain area (Liu Peiying et al. 2007).
4.1.2.1.2.2 ‘Yumo 1’ ‘Yumo 1’ is a group of naturally mutated individual plants optimized

by Southwest University from the cultivated groups of the A. konjac in Yunnan. It is an
excellent variety screened out by years of repeated selection, tissue culture and rapid
propagation, variety comparison tests, regional tests, and production tests. In 2001, tissue
culture, rapid propagation, and repeated selection were conducted on the mutated selected
individual plants; then, after systematic breeding of individual plants, the strains with
tall plants, dark green leaves, petioles with a ground colour of green and black spots, and
nearly circular corm were selected for a multiyear variety test. This variety was approved
by the Chongqing Crop Varieties Examination and Approval Committee in 2008 (Niu Yi
et al. 2010).
The mountainous areas with an altitude of 600–1,400 m in Sichuan, Chongqing, Yunnan,
and Guizhou are suitable for planting ‘Yumo 1’. This variety has a strong growth vigour
and green trisected leaves. The split leaves are pinnately divided or bipinnatifid. The small
split leaves are oblong and sharp pointed. The stalk has a ground colour of dark green
with black massive stripes. The corm is nearly circular and has a yellowish-brown epider-
mis. The interior tissues of the corm are white. The offset tubers naturally separate from
the parent upon maturity and shrivel into a garlic head-like shape. It takes about 130 days
from the emergence to maturity and senescence. The variety has a strong growth vigour
and stout plants. The yield per mu (1 ha = 15 mu) is 2,100 kg; the dry matter content in the
fresh konjac is 21.5%–22.2%; and the content of the KGM is 59.8%–60.2% dry basis. This vari-
ety is of high quality. The incidence of soft rot and southern blight is lower than in native
varieties of the A. konjac in Chongqing.
4.1.2.1.2.3 ‘Qingjiang A. konjac’ ‘Qingjiang A. konjac’ came from the systematic breed-
ing of the local genetic resources of the konjac in the Wuling mountainous area by the
Enshi Prefecture Academy of Agricultural Sciences and was approved by the Hubei Crop
Varieties Examination and Approval Committee. The seedling of this variety has a strong
growth vigour, uniform emergence, and palmately compound leaves with green or dark
green small split leaves; the petioles are stout with stripes; the corm is spherical with
brown epidermis; the bulbil is tip shaped and pink; and the interior is white. The whole
growth period of this variety is about 125 days. It is resistant to southern blight and soft
rot; its yield per mu is about 2,000 kg; the dry matter content in the fresh konjac is around
17.4%; and the content of the KGM is 51.4% dry basis. This variety is suitable for planting at
an altitude of 900 ~ 1,400 m in southwest China and is the dominant variety in the devel-
opment of the konjac industry in Enshi Prefecture and the Wuling mountainous area (Liu
Jinlong et al. 2010).
4.1.2.1.2.4 ‘Yunyu 1’ ‘Yunyu 1’ was bred from the high-quality genetic resource groups
of the Lijiang A. konjac in high attitude areas by the Yunnan Academy of Agricultural
Sciences and was registered with the Yunnan Provincial Horticultural Plant Variety
Registration Office in 2009. For this variety, it takes about 180 days from the emergence to
maturity and senescence. It is a medium-late maturing variety; the emergence is uniform;
and the seedling has a strong growth vigour. The leaves are green and trisected. The split
leaves are pinnately divided or bipinnatifid, or pinnately divided after dichotomous
branching. The small split leaves are oblong with gradually sharp points; the whole leaf is
large in area; and the plant is stout. The petiole is light green in the ground colour and has
greenish-brown spots, as the plant grows, the ground colour darkens to dark green and
the greenish-brown spots connect into one, making the whole petiole greenish-brown.
The corm of ‘Yunyu 1’ is oblate with few offset tubers; the hilar scar is smooth without
protuberance; and the interior tissue of the corm is white. The dry matter content in the
fresh konjac is 17.5%–20.3% and the content of the KGM 61.8% dry basis. This variety is of
high quality. The average yield is 37,500 kg/ha; the disease resistance is superior to most
cultivated varieties; and the incidence of soft rot and southern blight is lower than that of
the control variety. This variety is suitable for planting in the Yunnan-Guizhou plateau
areas with an altitude of 1,500–2,300 m (Li Yongjun et al. 2010).
4.1.2.1.2.5 ‘Qinmo 1’ ‘Qinmo 1’ was bred from the genetic resources of the A. konjac in
Qin-Ba Mountain areas by the Shaanxi Ankang Qin-Ba Konjac R & D Center. In 2002,
some naturally mutated individual plants with a compact plant architecture and strong
tolerance of shade were discovered among the cultivated plants of the A. konjac in Langao
County, Shaanxi Province. The corms were selected as parent material for propagation
and screening to select strains with uniform growth, strong resistance to diseases, few
offset tubers, regular corm, and high expansion coefficient. Then, after years of propaga-
tion, multipoint regional tests, and production tests, this variety was named as ‘Qinmo 1’
within the scheme of identification of nonmajor crop varieties in the Shaanxi Province (Li
Chuan et al. 2010).
The whole growth period of this variety is 160 days; the leaves are green or dark green
and trisected; the split leaves are pinnately divided or bipinnatifid, or pinnately divided
after dichotomous branching; the small split leaves are ovate triangular with gradually
sharp points; the plant architecture is compact with an included angle of 151.20° between
the bifurcation and the petiole; the petiole is dark brown with a few light pink flaky spots;
the corm is nearly circular and has a drab epidermis; and the interior tissues of the corm is
milk white. ‘Qinmo 1’ has a strong tolerance of shade and is suitable for underwood plant-
ing; the average field yield is 1,987 kg/mu; and the dry matter content in the fresh konjac is
21.21% and the content of the dry basis KGM is 57.77%. This variety is suitable for planting
in the Qin-Ba Mountain areas with an altitude of 700–1,200 m.
4.1.2.1.2.6 ‘Chumohua 1’ ‘Chumohua 1’ is a new variety of the A. konjac bred from the
local resources of the A. konjac by the Yunnan Province Chuxiong Prefecture Agricultural
Sciences Research and Promotion Institute and is characterized by a high yield, strong
resistance to diseases, and good processing quality. It was registered by the identification
of the Yunnan Seed Control Station. ‘Chumohua 1’ has a stable performance in agronomic
terms in different regions and a strong adaptability. It is superior to main local varieties
in terms of disease resistance, unit area yield, and the expansion coefficient of the corm
(Zhang Yan et al. 2013).
4.1.2.1.3 A. muelleri
A. muelleri refers to the konjac varieties that can grow bulbils on the leaf (similarly to
A. bulbifer). The A. muelleri has tall plants and strong heat resistance and is mainly dis-
tributed in the tropical rain forest in Southeast Asia and in southern Yunnan, China. The
A. muelleri has particularly rich morphological characteristics with significant differences
in petiole colour and stripes. The corm of the A. muelleri is usually oblate without offset
tuber. The A. albus and A. konjac grow fibrous roots at the shoulder of the corm only, but
the corm of the A. muelleri is covered with fibrous roots. The developed root system may be
the important reason for the tall plant, strong growth vigour, and rapid expansion of the
corm of the A. muelleri. The corm section is pink, yellow, or light yellow. The content of
the KGM in the corm is comparable to that in the A. albus and A. konjac. After flowering,
the A. muelleri can produce seed by ‘parthenogenesis’, which is one of the reasons why the
A. muelleri has attracted much attention. In recent years, influenced by market demand,
China has gradually imported the bulbils and seeds of the A. muelleri from Southeast
Asian countries for cultivation tests. At present, the approved or registered varieties of the
A. muelleri include ‘Mile 1’, ‘Mile 2’, ‘Mile 3’, and ‘A. muelleri’. Currently, the cultivated area
of the A. muelleri in Yunnan and Sichuan is about 10,000 mu. The successful introduction
and cultivation have greatly improved farmers’ enthusiasm for planting.
4.1.2.1.4 Yellow Konjac

The meat of the corm of yellow konjac groups is yellow, thus, such groups are collectively
referred to as yellow konjac. Actually, yellow konjac includes several different varieties.
Among these varieties, the A. ximengensis is of the KGM type and has a high KGM content.
A. yuloensis is of medium type and has a KGM content of about 33%; A. paeoniifolius and
A. dunnii distributed in Southern Yunnan, have no or little KGM, and are cultivated as
vegetables for food and as starch crops. At present, the yellow konjac groups are mainly
utilized as original resources. A few groups are used as parents in crossbreeding. Now,
the yellow konjac has been artificially cultivated in a small amount, thus, the potential of
variety breeding is huge.
4.1.2.1.5 Distant Hybrid Variety ‘Emoyu 1’

This variety is a distant hybrid variety of the A. konjac and A. albus. It is the seedling
obtained from the interspecific cross between the A. konjac ‘06–25’ variety from the Wuling
mountainous area (running from the Chongqing Municipality and East Guizhou to West
Hunan) as the female parent and the A. albus ‘2005Y-007’ variety from Yongshan County,
Yunnan, as the male parent. The individual plants that performed well were selected
from the F1-generation seedling for strain breeding. ‘Emoyu 1’, the new variety from sub-
sequent purification breeding and interspecies cross, was approved by the Hubei Crop
Varieties Examination and Approval Committee. The 2-year-old plants of this variety are
funnel shaped in plant architecture, with a plant height of around 110 cm, and an expan-
sion degree of about 100 cm. The petiole is green in the ground colour and covered with
green or dark green spots; and the leaflets are great in number and are long and elliptic.
The flower bud has a median size and coiled bud edge; and the appendage is dark purple.
The average yield per mu is around 2,500 kg. The corm column is spherical; the middle-
lower part is smooth; the shoulder is uneven; the epidermis is brown; the offset tubers can
be as many as 20 and are 4–5 times of those of ordinary A. konjac; and the corm is white in
meat and has a KGM content of about 55.85% dry basis. It is resistant to soft rot and grey
mould (Liu Jinlong et al. 2010).
4.1.2.2 Cultivated Species and Varieties in Japan

About 1500 years ago, the konjac was introduced from China into Japan along with
Buddhism. Then it gradually spread from the upper class into the folk, became a popular
food, and triggered the R&D of the konjac in Japan. In Japan, only the A. konjac Koch is
used in production. There are in total six varieties in Japan.
4.1.2.2.1 Native Variety

The native variety in Japan is called ‘Dayu variety’, ‘Shiyu variety’, or ‘Nanyang variety’
(Wakabayasi 1957). Tsuji (1990) mentions it as ‘endemic species’, the Japanese population
call it ‘Hongjing’, ‘Baiman’, ‘Ganyu’, ‘Heyu’, ‘Dayu’, etc. Now it is collectively referred to as
the native variety (Tsuji 1990). The native variety grown in different regions shows many
variations in its characteristics and disease resistance. However, it is difficult to divide the
specimens into different strains. In most cases, the plant architecture of the native variety
is horizontal. In some cases, the plant architecture is of semi-erect type due to different
climate or cultivation conditions. The whole leaf is large in area and light green; the peti-
ole is scattered with small stripes; the corm is slightly flattened; the epidermis has a dark
colour and many grooves; the bud eye is deep; the main bud is red; and the lateral corms
are very few and are longish corms with many stripes on the epidermis. This variety is
characterized by early maturity and large corm. The fine granules’ proportion in the flour
and the viscosity are high. It is of high quality, but has a weak resistance to leaf spot dis-
ease, root rot, and is prone to zinc and magnesium deficiency. Low temperature may eas-
ily cause yellowing, while high temperature causes sunscald. It is vulnerable to diseases
and natural calamities. Therefore, it is a variety that is the most difficult to plant. It is only
suitable for planting on mountain slopes without a sharp rise of temperature in the mid-
summer and with short sunshine duration.
4.1.2.2.2 Bittyu Variety

The okayama-ken variety, a representative strain, has semi-erect to horizontal plant archi-
tecture; and its leaflets are of large, long leaf type, thus, the leaf area is small. The corm is
in a shape similar to that of the native variety. The young corm is in a long cylinder shape.
The bud nest is shallow; the main bud is light red; and the traces of the offset tubers are
slightly raised points and are concentrated on the shoulder of the corm, which is different
from the native variety. The lateral corms are more spherical than in the native variety,
have few nodular stripes, belong to mid-maturation varieties, and have a relatively earlier
budding stage, leaf expansion stage, and good corm expansion. However, the expansion
is significantly reduced in the perennial corm. The content of the KGM in the corm is low;
the number of lateral corms is considerable; and its fertility is higher than that of the native
species. It has stronger resistance to diseases and natural calamities than the native species
and is suitable for cultivation in the regions with the same conditions as the native variety
or regions with slightly higher temperatures and slightly stronger sunlight. Since it has a
low yield and inferior quality, it is often cultivated in marginal areas of the land suitable
for the konjac (GPAIA 1991).
4.1.2.2.3 Chinese Variety

At a point in time, a well performing konjac variety was introduced from China for cul-
tivation. The plant architecture is of erect type; the leaflets are of small broadleaf type,
are great in number, and are dark in colour; and there are small protuberances and small
white spots on the surface of the petiole, thus, it is not as smooth as the native variety.
Since the stripes are large and interconnected, the colour is darker, and the whole petiole
appears black. The corm is nearly spherical with few grooves on the surface; the colour of
the epidermis is light; and the bud eye is shallow. The offset tubers are light brown and
concentrated on the middle-upper part of the corm, which is long-shaped with a slightly
larger upper part, and no separation layer is formed. This variety is late-maturing with
a late budding period, but the time from budding to leaf expansion is short. The leaf
expansion period is similar to that of the native variety. The expansion ability of the
corm is considerable. The dry matter content is very low, and the coarse particle propor-
tion is 8.3%. However, the KGM content of the powder processed from the dry konjac
pieces can be up to 65.5%; and the viscosity of the fine powder is also high. The plants
have many offset tubers and strong resistance to such natural hazards as high tempera-
tures, strong sunlight, too wet or too dry soil, and wind damage, also leaf blight and root
rot. Therefore, this variety can be regarded as robust. However, once a plant is infected
with soft rot, the progression of the disease and the infection of the surrounding healthy
plants are very rapid, the corms are rotting during growth, storage, or drying. Since this
variety is late-maturing, it can grow and develop even in late autumn and thus allows
the corm to fully expand. Therefore, it is suitable for planting at a low altitude. However,
the plants cannot fully grow and develop on hilly land with early frost, as it leads to a
low yield and quality and poor storage performance of the corm. In order to prevent rot
disease, the best choice for planting this variety are areas with few natural hazards, such
as hail or a typhoon.
4.1.2.2.4 Haruna-kuro (Nonglin 1)

It is a variety derived from the interspecific hybridization between different species of
the A. konjac. It was bred in 1949 and was the seedling offspring of the F1 generation from
the artificial hybridization with the varieties from China as the female parent and the
native variety as the male parent conducted by the Japanese Agriculture, Forestry and
Fisheries in 1948. The yield and traits identification tests were conducted with the system
name of ‘Qunxi 1’ in 1959. It was registered as ‘Konjac Nonglin 1’ and was named Haruna-
kuro. The name comes from the famous mountain ‘Haruna’, where this variety is grown
in Gunma Prefecture and also indicates that its petiole is black. The plant is ‘Y’ shaped;
the leaflets are of slightly larger broadleaf type, dark green in colour and glossy. There are
small protuberances and small white spots on the surface of the petiole. However, there
are less of them than on the variety from China. The petiole is light red in ground colour,
but the black stripes on the petiole are large and continuous, making the whole petiole
look black. The morphology of the aboveground part is like its female parent, the species
from China, so that it is difficult to distinguish them. Nevertheless, this variety has larger
leaflets, better gloss of leaves, and fewer small white spots on the petiole. The corm has a
slightly flattened spherical shape, which is similar to the native species. The corm is more
spherical as it grows. There are fewer grooves on the surface. The epidermis is light brown;
the bud eye is deep; and the main bud is bright red. The upper part of the section of the
corm is light red, KGM crystals can be seen, the offset tubers are large and equivalent to
the native species in number, and can be easily separated from the corm during harvest-
ing. The expansion of the corm is excellent and is considered intermediate between the
variety from China and the native varieties. The konjac flour yield is lower than that of the
native varieties, but higher than that of the species from China. The proportion of large
particles and the viscosity of the fine particles are higher than that of its parents. The ger-
mination and leaf expansion of this variety occur early; and its maturation period is close
to that of the native varieties, thus, it is a mid-maturation variety. Its resistance to sunscald,
yellowing, and other climatic hazards is stronger than that of the native varieties, while
its resistance to soft rot and dry rot is similar to that of the native varieties. However, its
resistance to leaf blight and root rot is weak, thus, its yield will not exceed 30% of that of
the native varieties, which is its demerit. Since its maturation period is slightly late, it is
suitable for planting in the moderately hilly areas with an altitude of 200–400 m in Japan
(Yamaga Ichiro 1966).
4.1.2.2.5 Akagi-ohdama (Nonglin 2)

It is a new variety bred from the intraspecific cross between different varieties of
the A. konjac. The artificial crossbreeding was conducted in the Gunma Prefecture
Agriculture Proving Ground in 1953 with the species from China as the female parent
and the native species as the male parent. It was registered as ‘Konjac Nonglin 2’ in
1970 and named Akagi-ohdama (あかぎおおだま). This variety was named after the
‘Akagio’ in the Gunma Prefecture, where it was bred and first popularized. It has an
outstanding expansion of the corm (‘dama’ means large corm in Japanese). This variety
is characterized by a horizontal ‘T’-shaped plant architecture and a blade longer than the
petiole. The split leaves are of broadleaf type and large in number. The leaf colour is the
lightest among the existing varieties in Japan. The petiole has a light red ground colour,
with small protuberances, and many small white spots on the surface. The stripes are
medium in size and are distributed continuously, but lighter in colour than the variety
from China. The old corm is slightly oblate and rounder than the variety from China.
The young corm is irregular in shape and looks like the specimens from China. The epi-
dermis is brown; the bud eye is deep; and the main bud is deep red. The lateral corms
grow on young corm. A few lateral corms can be spherical, but generally have an elon-
gated shape. Like the variety from China, the epidermis is smooth and light in colour;
and the separation from the konjac flour during processing is poor. The storage loss is
huge; thus, postharvest operations must be conducted immediately after harvesting
to preserve the corm quality. This variety has more lateral corms than the native vari-
ety; the single weight is high; and the propagation rate is good. The variety has a high
expansion of the corm and high yield. The proportion of coarse particles in the konjac
flour is low and, thus, the ratio of fine/coarse particles is high, which is superior to
the native variety. The flour is characterized by high viscosity. Like the native variety
and Haruna-kuro, this variety is a mid-maturation variety, but its germination and leaf
expansion are early. Its resistance to sunscald and deficiency of elements is stronger
than that of the native variety, but weaker than that of the variety from China. Its resis-
tance to yellowing and low temperature damage is strong, but the resistance to soft rot
is weak; the resistance to leaf blight is strong; and that to root rot is between the native
variety and the variety from China. It can be planted in the medium and high hilly areas
with an altitude of 300–600 m. For planting in the mountainous areas with an altitude
of around 600 m, the plastic film mulching can ensure safety. It should not be planted
in the plots prone to sunscald at low altitudes, high temperatures, and strong sunlight.
It is suitable for planting in the east mountain areas of Kanto (Honshu Island). Since its
corm has good expansion and many lateral corms, 2-year-old corms should be used for
cultivation (Yamaga Ichiro 1970).
4.1.2.2.6 Myogi-yutaka (Nonglin 3)

Myogi-yutaka was bred, with the objectives of high yield and high quality, from the
offspring of the hybrid plants. Breeding work was carried out in the Gunma Prefecture
Agriculture Proving Ground with ‘Qunxi 26’ as the female parent and a high yield strain
of the variety from China as the male parent in 1971. A preliminary production perfor-
mance test was first conducted in 1981. The characterization test was carried out with the
local name of ‘Qunxi 66’, and the breeding ended in 1991. It was registered as a new variety
with the name of ‘Konjac Nonglin 3’ and named Myogi-yutaka (みょうぎゆたか). Its name
came from the name of Mount ‘Myogi’ and the meaning of high yield. The multi-point
cultivation test on the adaptability of this variety had been carried out in various regions
since 2001. After the introduction and cultivation by Nagano-ken, Saitama-ken was the
second prefecture to introduce and cultivate this variety. This variety has strong resistance
to leaf blight and root rot, which is equivalent to the variety from China. The yield of fine
particles in the flour is 20% higher than that of Haruna-kuro, 10% higher than that of the
variety from China, and also much higher than that of Akagi-ohdama and other varieties,
but equivalent to that of the native variety. The viscosity of the fine powder is also higher
than that of any other varieties. The Myogi-yutaka is suitable for planting in warm flat
plots with an altitude of 100 m and middle altitude mountainous areas in the Chichibu
District, Saitama Prefecture (Honshu Island).
According to the statistics of the Japan Gunma Prefecture Comprehensive Agriculture

Proving Ground, leaf blight is the most serious disease among the konjac diseases in Japan.
Among the six abovementioned varieties grown in Japan, the incidence of leaf blight in the
native variety is high. According to the records, its incidence was up to 100% in September
1983. In addition, the sunscald is also more serious than in other varieties. However, the
coarse particles and fine particles proportions are higher in contrast to other varieties. Since
the expansion ability of the corm of the Bicchu variety is low and the corms are small, the
yield per surface area, fine particles rate (relative to coarse particles), and the viscosity are
low. The corms of the variety from China have a higher yield. Although the incidence of
leaf blight and that of sunscald is low, its resistance to bacterial soft rot is particularly weak.
In spite of the fine particle proportion and viscosity being good, its coarse particles propor-
tion is low. The cultivated area of the aforementioned three varieties (native, Bittyu, and
Chinese) has gradually decreased, and they became replaced with Haruna-kuro, Akagi-
ohdama, and Myogi-yutaka produced by crossbreeding. The expansion ability of the corm
of the Haruna-kuro is the highest in the first year and becomes medium in the second
and third year. The coarse particles proportion of the Haruna-kuro is high and the fine
particles proportion moderate, but the viscosity of the fine particles is high. Its resistance
to leaf blight and root rot is weak. The weight and the expansion ability of the corm of
the Akagiodama in each year are high and the yield is high. Although its coarse particles
proportion is low, its fine particles proportion is high, and the fine particles proportion of
3-year-old and 2-year-old corms can be higher than 68%. The viscosity of the fine particles is
also high. Its resistance to leaf blight is strong; its resistance to root rot is between that of its
parents, but its resistance to soft rot is weak; and sunscald can easily be caused under high
temperature and drought conditions. However, its resistance to yellowing and other low
temperature damages is strong, and, thus, it can be planted at high altitudes.
The trend of selecting varieties throughout Japan is as follows: the Bittyu variety is being
replaced, the cultivated area of the native variety is extremely small, and that of the variety
from China is also decreasing year by year. In contrast, the area under the Haruna-kuro
is increasing, the cultivated area of the Akagi-ohdama is currently the largest, and the
Myogi-yutaka is slowly taking the shares of other varieties in virtue of its many merits.
The overall proportion of varieties in Japan in 1999 was approximately as follows: Haruna-
kuro 20%, Akagi-ohdama 62%, the native variety 9%, the variety from China 2%, with
Bittyu and Myogi-yutaka accounting for the remaining 7%. The variety structure in the
Gunma Prefecture was: the native variety 0.3%, the variety from China 3.7%, the Haruna-
kuro 28%, and the Akagi-ohdama 68%. In 1999, the harvested area in Japan was 3,484 ha,
Gunma Prefecture 2,540 ha, the total national yield was 62,600 t, and Gunma Prefecture
53,600 t. The yield of the Gunma Prefecture appears to be correlated with the variety struc-
ture (Japan Konjac News Agency 2001).
4.1.2.3 Cultivated Species and Varieties in Other Countries and Regions

4.1.2.3.1 Southeast Asia
Southeast Asia includes Burma, Laos, Vietnam, Thailand, Indonesia, and Malaysia and
is the centre of origin for the konjac, thus, the konjac is widely distributed in Southeast
Asia. The growing season for the Amorphophallus species in Southeast Asia is the rainy
season. The A. muelleri and yellow konjac groups have high economic value and, thus, are
the mostly harvested species. In the past 10 years, influenced by Chinese market demand,
the cultivation of the konjac in Southeast Asian countries has been developed gradually
(Nikmah et al. 2016; Yuzammi et al. 2017; Lontoh et al. 2019).
4.1.2.3.2 India
Some 13 varieties of the konjac are growing in India, but only starch type, the A. paeoniifo-
lius, is cultivated and consumed at a large scale. The main cultivated variety is the Kavvor.
The other variety is the A. paeoniifolius var. hortensis Backer. According to Anil et al. (2011),
the specimens of 12 varieties of the A. paeoniifolius collected in Tamil Nadu showed high
variation: the yield of a single corm 100–3,000 g and the starch content 14%–36%. The
A. paeoniifolius is consumed in India as a vegetable with the corm washed, sliced, and cooked.
4.1.2.3.3 Other Regions

Various species of the Amorphophallus are distributed in the centre of origin of this genus,
Indochina and Africa. However, there is no significant cultivation and consumption of the
konjac in Africa.
4.1.3 Elite Breeding of Konjac

In cultivation, the konjac is a perennial asexually reproduced crop that can be harvested
2–3 years later. The annual reproduction coefficient of potatoes, sweet potatoes, and other
tuber crops can be up to more than a dozen times, but that of the konjac is only about
4–8 times. In addition, potatoes, sweet potatoes and other tuber crops can be cut into pieces
for reproduction, but the konjac is seldom cut into pieces for reproduction for its apical domi-
nance is extremely obvious, its epidermis is thin and meat crisp, thus, it may easily be injured
and affected with soft rot. The reasons for the variety degeneration of the konjac mainly
include a decreased resistance to diseases (soft rot, southern blight, etc.), sensitivity to high
temperature, sensitivity to high light intensity, decrease in the number of offset tubers, and
inappropriate offset tuber to corm weight ratio. The large-scale and long-distance transit of
seed corms with nonstandard packaging is the primary reason for the failure in remote kon-
jac planting areas. An elite cultivar is a high yielding cultivar possessing various desirable
traits. These can be used in crosses with other elite lines, and the breeding can focus on the
trait of interest, as there is minimal disruption of the elite genetic background. The commer-
cialization of elite konjac varieties and the local production of the commercial seed corms are
effective measures to improve the benefits of the konjac planting.
4.1.3.1 Establishment of Elite Variety Breeding Grounds for Konjac

The konjac has a large range of suitable planting areas. In industrial development, in order
to prevent damage to seed corms from diseases due to long-distance transportation, pro-
fessional elite breeding grounds for the konjac can be established by districts in combina-
tion with the suitable areas of the konjac cultivation and according to the needs of the
konjac industry. The following sections describe the principles of elite breeding in China,
but could apply to other konjac producing countries. The main areas suitable for produc-
ing the konjac in China and for the establishment of the elite breeding grounds have been
identified as follows: Yunnan-Guizhou Plateau, Wumeng Mountains area (Yunnan), moun-
tain areas around Sichuan Basin, Wuling Mountains area (Chongqing Municipality and
East Guizhou to West Hunan), and Qinba Mountains area (southern Shaanxi Province).
The breeding of the elite konjac varieties should include the introduction of performing
new varieties and the matching cultivation techniques and should be gradually developed
from small to large scale to progressively implement the local production of commercial
seed material. On this basis, the konjac test demonstration bases can be built for variety
improvement and the demonstration of cultivation techniques.
4.1.3.2 Principles of Elite Breeding System for Konjac

Since the epidermis of the konjac is thin and the amount of seed corms is considerable,
large-scale and long-distance transportation of seed corms with nonstandard packaging
makes seed material prone to scars before being planted and is the primary cause for dis-
ease prevalence and planting failure. Therefore, the solution to the problem of industrial
development must be based on the establishment of local elite breeding systems. The elite
breeding system for the konjac includes: three-stage breeding system, 2-year rapid breed-
ing technology for the test-tube plantlet (test-tube konjac), and 2-year rapid growth tech-
nique for offset tubers.
4.1.3.2.1 Three-Stage Breeding System for Konjac

The three-stage breeding system for the konjac mainly refers to the establishment
of the ‘breeder’s seed—foundation seed—elite seed’ three-stage breeding ground.
The elite breeding ground for each stage should be established in areas with suitable
temperatures, fertile soil, loose soil, and good shade conditions for convenient produc-
tion and supply.
The breeding grounds for the breeder’s seed are usually established in plots where no
konjac is going to be commercially grown within 3 years. The seed corms and 2-year-old
offset tubers of the robust konjac plants with marked variety characteristics, high yield,
and strong resistance to diseases (mainly soft rot and southern blight) are selected. In addi-
tion, the corms for reproduction should be regular in shape with a shallow bud eye and
without wounds, disease, or insect presence. The breeding grounds for the breeder’s seed
usually are established at the prefecture-level agricultural technology promotion stations
or research institutes with professionals who follow up and manage the field operation
throughout the breeding process.
The breeding grounds for the foundation seed are usually established in plots where
no konjac is going to be commercially grown within 2 years. The corms and offset tubers
of the plants of the breeder’s seed are used for reproduction. In the process of produc-
tion of the foundation seed, the latter should be scrutinized for signs of variety degenera-
tion. Only 1-year-old and 2-year-old corms and offset tubers without signs of degeneration
should be chosen for planting on the elite breeding grounds. The breeding grounds for
the foundation seed should be established at a county level. The management of the field
operations should be done by trained professionals.
The foundation seed for reproduction on the elite breeding grounds is selected on the
basis of age and weight. The plots where no konjac is going to be grown commercially
within 2 years are usually selected for reproduction. The grounds for the elite seed should
be established at the township-level agricultural technology promotion stations in main
producing areas. The elite seed should be supplied locally (Xiang Changqing et al. 2011).
4.1.3.2.2 Two-Year Rapid Breeding Technology for Test-Tube Plantlet

The test-tube plantlet of the konjac (test-tube konjac) refers to the single test-tube plantlet
produced in a tissue culture vessel or small corms directly produced by tissue culture in
a test tube by a specialized factory. This method ensures rapid reproduction, a 10–30 g
small corm of the konjac can reproduce hundreds of thousands or even millions of
2–30 g breeder’s seed for planting on dozens of mu in a greenhouse or seed konjac breed-
ing ground. This technique enables the promotion of the new konjac varieties with a high
yield, high quality, and high disease resistance within a relatively short period of time.
Also, the meristem culture can be used to produce decontaminated konjac breeder’s seed.
4.1.3.2.3 Two-Year Rapid Growth Technique for Offset Tubers

In the first year, the stout and well-developed offset tubers with a full terminal bud and
without disease or wound are selected. They are subjected to accelerated germination,
reasonably closely planted, and carefully managed. The weight of offset tubers can be
increased by 10–15 times to become 150–200 g small corms in the same year. In the sec-
ond year, after reasonable planting, the abovementioned small corms can grow into about
1,000 g konjac corm with 5–8 offset tubers. Thus, the traditional 3-year planting of the com-
mercial size konjac is shortened to a 2-year planting.
4.1.3.3 Use of Sterilized Konjac Seed Material

Since the corm is the reproductive organ of the konjac, viruses can accumulate in seed
corms from generation to generation, causing multiple morphological abnormalities
from leaf expansion, such as insufficient expansion of plants and reduced photosyn-
thetic efficiency. Such abnormalities may cause a decline in the yield and quality dete-
rioration, thus leading to variety degeneration. At present, the konjac production is
bothered by soft rot and southern blight and, thus, much attention has been paid to these
two diseases. However, from the perspective of the breeding of quality konjac seed,
importance should also be attached to the decontamination of the konjac. The varieties
or strains that are cultivated in various regions and have a high yield and good quality
are often used as the reproduction materials in the meristem sterilization technique for
tissue culture of the konjac following the sterilization procedure used in the seed potato
production. The test-tube plantlets for tissue culture are strictly tested and reproduced
in an enclosure protected from insects with a 40-mesh net for more than 2 years after
virus detection. During that time, the morphology, quality, and productivity of plantlets
are tested. The high-quality decontaminated breeder’s seed can be obtained by select-
ing from several asexual strains. The quality of the decontaminated konjac seed will be
tested and virus checks performed by seed inspection authorities whenever possible
(Dai Qingtang et al. 2011).
4.1.3.4 Local Production and Commercialization of Elite Varieties

There have been numerous failures of the konjac cultivation caused by the long-distance
transportation in the development of the konjac industry in various regions. The konjac
corms have high water content, with thinner epidermis, and more crisp meat than the
corms of other tuber crops. As a result, the konjac corms are more likely to be injured.
The endemic soft rot caused by the injuries to the seed corms, which are caused by
the long-distance transportation of large amounts of the seed konjac without appropri-
ate packaging, is the primary cause for plant failure. Therefore, the planting material
should be produced on the premises to ensure sufficient supply of the konjac seed to
the growers.
As corms and offset tubers are generally used for the traditional konjac planting, the
konjac is a typical asexually reproduced crop. Therefore, as with other asexually repro-
duced crops, it is difficult to identify and distinguish individual plants of a different vari-
ety in a plantation. However, in recent years, with the growing market demand for the
KGM, the demand for high-quality konjac seed has increased. Given the limitations to
the development konjac plantations caused by long-distance transportation of seed mate-
rial, it is necessary to implement standardized production, packaging, and distribution
of the elite seed konjac near the plantation sites. The required measures should include
the promulgation of national or industrial standards for the elite konjac varieties, the cre-
ation of brands for the high-quality konjac varieties, and the standardization of produc-
tion, packaging, and transportation of the elite varieties. Positioning the production of the
seed material near the plantations will facilitate the commercialization of the elite konjac
varieties and reduce the cost of the seed material.
4.1.4 Ecological Adaptability of Konjac

The konjac is an undergrowth plant from the Southeast Asian rainforests. The original
environment gives unique ecological habits to the konjac. Although the current konjac
varieties for cultivation and production are spread from south to north, after thousands
of years of planting and domestication by mankind, they still follow many original habits,
such as growing well in shaded, warm, and moist conditions and avoiding high or very
low temperatures and bright light. Meanwhile, the konjac is similar to other tuber crops in
that the expansion of its underground corms has particular requirements regarding soil
permeability, soil quality, and also water and fertilizer supply.
4.1.4.1 Influence of Temperature on Growth and Development of Konjac

4.1.4.1.1 Influence of Temperature on the Growth of Different Parts of Konjac
The konjac corms begin to germinate around 5°C. The optimum temperature for germi-
nation is 20°C–25°C. If the temperature is higher than 45°C or lower than 0°C, the bud
dies after about five days. The bud grows slowly under 15°C; and it grows rapidly above
35°C, but the bud is delicate and the leaves are curled upon emergence. The konjac
leaves grow normally between 20°C and 30°C. At such temperatures, the chlorophyll
content is the highest and the photosynthesis maximum (Zhang Xingguo et al. 1991).
The injury of the konjac leaves due to temperature is reflected in the changes in the cell
membrane permeability. Within the range of 15°C–35°C, the cell membrane permeabil-
ity is constant. When the temperature is lower than 15°C, leaves gradually turn yellow
and start wilting. Above 35°C, the leaves slightly fade for the chlorophyll is degraded.
The wilting and sagging of the leaves appear clearly at a sustained temperature above
40°C, showing obvious signs of injury (Zhang Xingguo et al. 1992). The optimum tem-
perature for the development of the root system of the konjac is 20°C–26°C. The root
system stops growing under 5°C and above 35°C. The optimum temperature for the
development of the konjac corms is 22°C–30°C. A large temperature difference between
day and night promotes the accumulation of dry matter, resulting in an increased yield
and better quality. The konjac corms are not resistant to low temperatures. The intra-
cellular water freezes below 0°C, damaging the cell structure and making the whole
corm inactive. Therefore, the thickness of the frozen topsoil layer should be taken into
account for planting the konjac in areas at high altitudes and high latitudes. In areas
with freezing topsoil, the konjac cannot survive the winter outdoors. Thus, corms must
be protected for overwintering by planting them deeper or digging them out and stor-
ing them indoors at subzero temperatures during the winter and replanting them in the
spring (Yang Xiulian et al. 2010).
In conclusion, the konjac is a crop that likes a warm environment, but with temperatures
not higher than 35°C and is sensitive to temperature changes. The temperature directly
influences the growth, development, cultivation, and production of the konjac. The tem-
perature suitable for the growth of the konjac is 5°C–35°C; and the optimum temperature
is 20°C–25°C in the growing season of the konjac.
4.1.4.1.2 Influence of Accumulated Temperature on Konjac

The konjac can start its growth at temperatures above 5°C and needs a certain total tem-
perature to complete its life cycle. The A. albus can be planted in low mountain areas with
an altitude of 800 m and needs an active accumulated temperature of 4,863.1°C from ger-
mination to the senescence of the leaf. The effective accumulated temperature is 1,658.1°C.
The A. konjac grows well in mountainous areas at an altitude of 800 ~ 2,500 m and needs
an active accumulated temperature of 4,279.8°C and an effective accumulated temperature
of 1,089.3°C. Such accumulated temperatures determine the geographical distribution of
the konjac and are the important factors that should be taken into account when planning
the introduction and cultivation of this crop.
4.1.4.1.3 Temperature Requirement of Different Amorphophallus Species
Different konjac species have different requirements for temperature. The A. paeoniifolius
and A. dunnii are distributed at low-altitude areas in southern Yunnan characterized by a
warmer climate than the higher altitude areas in the same region. They belong to the heat-
resistant Amorphophallus species. The A. albus is mostly found in the dry-hot valley of the
Jinsha River in Sichuan and Yunnan and is more heat-resistant than the A. konjac.
4.1.4.2 Influence of Light on Growth and Development of Konjac

The development of the konjac in the undergrowth of rainforests forms the character-
istics of the konjac as a semi-shade plant that likes scattered light and avoids high light
intensity. The konjac has different response to light in different growth periods: when leaf
expansion has just been completed at the early growth stage, the light saturation point of
the konjac is low. The period when leaves grow and get mature is the stage when the kon-
jac grows most vigorously, and its light saturation point is the highest, then it decreases
gradually with aging. The A. konjac usually reaches its maximum light saturation point
or illuminance of about 22,000 lx around mid-August. The light saturation point of the
konjac throughout the growing period is between 17,000 and 22,000 lx, thus, it is a typical
semi-shade plant. The light compensation point (compensation point is the light intensity
on the light curve where the rate of photosynthesis exactly matches the rate of cellular
respiration) remains mostly stable at around 2,000 lx. The light intensity can effectively
improve the photosynthetic rate of the konjac before reaching its light saturation point.
When the light intensity reaches the light saturation point of the konjac, not only does the
continuous high light intensity cause curled leaves of the konjac, but also the accompany-
ing high ambient temperatures of high lights will cause sunscald of the konjac, decreasing
the photosynthetic efficiency. Yet, although the konjac is a shade crop, the sunshine dura-
tion should also be 9–10 hours.
Since the light intensity is usually too high, upon cultivation, it cannot be fully ensured
that the light intensity meets the demand range of the light saturation point of the konjac.
Studies have shown that within a certain range, the weaker the light intensity, the higher
the content of chlorophyll; and moderate shading can significantly increase the chloro-
phyll content in the konjac leaves. In addition, compared to the open field monoculture
planting, the konjac plants that are provided with 75% shading with a sunshade net have
long and thick petioles, a higher seedling index, higher chlorophyll content, higher photo-
synthetic rate, and significantly higher yield and KGM contents.
The konjac likes scattered light and low light intensity. In a greenhouse, different
light quality can be obtained by covering the greenhouse with red, green, blue, pur-
ple, and white plastic films. The konjac plants growing under the red film showed an
obvious growth advantage: had long, thick petioles and a significantly higher seedling
index than the control or the konjac plants under other films. They also had the high-
est chlorophyll content and photosynthetic rate, a higher corm yield, and KGM content.
Moreover, the soluble protein content and the starch content in the konjac corms grow-
ing under the red film were significantly lower than of those covered with other films
(Li Nanlin 2014).
There also is a difference in the photosynthetic performance between different
Amorphophallus species. Compared with the A. albus, the A. konjac has a higher net photo-
synthetic efficiency during the same growth period. The A. muelleri, A. dunnii, and A. paeo-
niifolius have a higher resistance to high light intensity. Also, the level of required shade
density varies with the environment, higher shade densities (60%–90%) should be adopted
in areas with high temperatures and long and strong exposure to sunlight; and the shade
densities of 40%–60% should be adopted in areas with short exposure and weak sunlight;
no shade treatment is needed in areas with poor light. For instance, in mountain areas
where the konjac is in the shade of mountain chains and numerous trees, temperatures
are low and humidity high, shading can be generally avoided. The higher the latitude or
altitude, the lower shade density that is needed (Kobayashi 1964).
In conclusion, the intercropping of the konjac with corn or other long-stalk crops, plant-
ing under sparse forest, or the large-area planting under economic fruit trees can make
the light intensity and light quality suitable for the konjac and improve the plant growth
and konjac yield.
4.1.4.3 Influence of Moisture Conditions on Growth and Development of Konjac

The konjac likes a moist environment, avoids waterlogging, and is not drought-resistant.
The konjac needs a uniform and sufficient moisture supply in the growing season. Studies
have shown that the konjac needs a high humidity soil environment in the early growing
and corm expanding period, when the optimum soil water content is about 80%. However,
the soil water content should be appropriately controlled and decreased from 80% to 60%
in the late growing period after mid- and late September in order to promote the accu-
mulation of nutrients in the corm. In the winter, the konjac corms are in the dormancy
state and need relatively dry soil conditions. A short-term (about one week) drought in the
growing period has little influence on the growth of the konjac. However, the long-term
drought has great influence on the yield of the konjac by causing the water stress of the
konjac plants and curled and yellow leaves. Serious droughts are usually accompanied by
the combined stress of high light intensity and high temperature, resulting in the wilting
of plants and shrinkage of petioles. In particular, the water shortage at the early stage of
corm expansion, followed by sufficient water supply at a later stage can cause a fracture
of the epidermis of the corm and lead to bacterial invasion and disease outbreak, result-
ing in a substantial reduction in production (Zhang Xingguo et al. 1992). A too high soil
humidity is not favouring the konjac growth. Since the ventilation tubes in the fibrous
root of the konjac are narrow, in the case of excessive rainfall with water ponding in the
field and excessive soil humidity, the permeability is reduced. This is unfavourable for gas
exchange, causes poor breathing of the roots and corms, causes rot in the field or during
storage, and seriously affects the yield and quality. During cultivation, plots should be
selected according to the abovementioned characteristics of the konjac, i.e., liking moisture,
avoiding waterlogging, and being not drought resistant. Drainage, moisture conservation,
and drought prevention operations should be carried out according to the environmental
conditions and development of the plant.
4.1.4.4 Mineral Nutrition

The growth of the konjac plants, the accumulation of reserve substances, and the yield for-
mation are related to mineral nutrition. The konjac plants have similar demands for nitro-
gen, phosphorus, potassium, and other essential nutrients as do potatoes and other tuber
crops. They also have different demands for mineral nutrition in different growing stages.
The konjac absorbs mainly potassium, followed by nitrogen, and then phosphorus fertil-
izer throughout the growth period. From the germination of the seed konjac until emer-
gence, the nutrients are provided by the seed corms, thus the formation and emergence
of the new plant can be ensured under adequate moisture and temperature. However,
after the emergence and leaf expansion, especially when the new corm expands rapidly,
abundant and adequate mineral nutrition should be supplied. At that stage of corm devel-
opment, the demand for nitrogen, phosphorus, and potassium of the plant is much higher
than during the maturation period of the corm. With the ageing and senescence of the
leaves and petioles, some nutrients are transferred to the corm for storage.
The nitrogen can promote the growth of the aboveground part, make the leaf dark green,
increase the efficiency of light energy utilization, accelerate the accumulation of organic
substance, and promote the formation of proteins or enzymes. Excessive nitrogen fertil-
izer causes an excessive growth of the aboveground part, decreases the resistance to plant
diseases, and promotes insect attacks. In the case of drought, excessive nitrogen fertilizer
may easily cause the insufficient maturity of leaves, which is bad for water retention. In the
case of insufficient nitrogen fertilization, the konjac plants are thin with low chlorophyll
content, which greatly influences the growth and expansion of the corm. From the begin-
ning of the rapid growth of the konjac corm, the nitrogen absorption of the plant remains
at a high level. From the onset of ageing, the uptake of nitrogen decreases rapidly, and it is
transferred to the corm for storage.
Phosphorus is involved in the glucose metabolism and nucleic acid synthesis. Although
the konjac does not need much phosphorus, it is still an essential mineral element for the
konjac. A sufficient supply of phosphorus fertilizer not only promotes the normal develop-
ment of the konjac plants, but also improves the yield and quality of the corms.
Potassium can promote the synthesis of the photosynthetic products and the transfer of
such products to the corm, increase the content of the starch and KGM in the konjac, and
has obvious effects on the quality and stability in the storage of the corm. It can promote
the robust growth of the konjac plants and increase the disease resistance. Therefore,
from the onset of growth of the konjac corm, the supply of potassium is of vital importance
in the rapid expansion period of the new corm. Besides, the konjac is a chlorine sensitive
crop. Therefore, when applying potassium fertilizer externally, potassium sulphate is usu-
ally selected instead of potassium chloride (Gu Tao 2017).
Calcium is an element that forms the cell wall. In the case of calcium deficiency, the meri-
stem suffers first, the formation of the cell wall is blocked, and the cell division reduced.
An adequate calcium nutrition can improve the disease resistance of the konjac plants.
Calcium combines with oxalic acid to form insoluble calcium oxalate crystals that cannot
only neutralize the acid, but also promote the formation of KGM crystals.
Trace elements, such as copper, boron, and zinc, also have great impacts on the growth
and development of the konjac. As a cofactor of copper oxidase, copper can enhance res-
piration and increase the net photosynthetic rate, increase the content of chlorophyll and
proteins, delay leaf senescence, and improve the resistance of plants. Boron can combine
with free state sugar, promote the transportation of sugar, and increase the yield of the
corm (Niu Yi 2004).
Organic fertilizer cannot only increase the content of organic matter in soil, improve soil
structure, and increase soil permeability, but also provide a lot of mineral nutrients. Under
certain fertility conditions, the amount of organic fertilizer applied is closely correlated to
the konjac yield.
According to the growth characteristics of the konjac plants, the fertilization principle
is ‘focusing on the application of base fertilizer and applying topdressing timely’. Good
understanding of the nutritional needs of the crop and the timely application of fertilizers
are the important guarantees for improving the konjac yield and quality.
4.1.4.5 Soil
The konjac corm grows in the ground and, thus, has high requirements for soil condi-
tions. Light sandy loam soil with a deep soil layer, loose texture, rich organic matter, and
good ventilation and drainage is the most suitable bed for the growth of the konjac. Loose,
thick, and fertile soil is an important guarantee for the development of the root system
and corm expansion of the konjac. Sticky and heavy soil with poor drainage, permeability,
and easy hardening is unsuitable for the growth of the konjac. The konjac planted in such
soil not only is low in yield, easy to become infected by diseases, but also has corms in
irregular shape and coarse epidermis that are bad for the konjac corm processing. A too
shallow soil layer is unsuitable for the growth and expansion of the konjac corm (Cui Ming
et al. 2006).
Most konjac varieties are suitable for growing in the slightly acidic soil with a pH value
of about 6.5 and can also be planted in neutral and slightly alkaline soil, but too acidic or
alkaline soil is unsuitable for the growth of the konjac; and there are serious diseases in
acidic soil. Compared with the A. konjac, the A. albus prefers slightly alkaline soil. That is to
say, the yield is relatively higher when the pH value is 7.0–7.5 and is greatly reduced when
the pH value is below 6.0 or greater than 8.0.
Physical and chemical conditions of the woodland or vegetable garden soil in front and
behind the house are suitable for the expansion and growth of the konjac corms. When
the konjac cultivation is transferred to the field, the planting bed should be improved
accordingly by increasing soil layer thickness, improving soil structure, adding farmyard
manure to increase the organic matter in the soil, retain water, and adjust the pH value of
the soil to promote the reproduction of beneficial microorganisms in the soil.
4.1.4.6 Other Conditions

4.1.4.6.1 Wind
The konjac plants are composed of leaves and petioles; and petioles are the only support-
ing structure of the konjac plants, but break and bend easily. The konjac only grows one
compound leaf in a growing season. Once the compound leaf is damaged, the yield will
be greatly affected. Breezes help to improve the ecological conditions between plants and
promote the growth and substance accumulation of the konjac; and violent winds may eas-
ily cause lodging and breaking of the konjac plants. Therefore, in areas with typhoons and
gale winds, the konjac should be planted in sheltered plots to avoid damage.
4.1.4.6.2 Shade Crops

The konjac is often intercropped with long-stalk crops, such as corn, sorghum, or sun-
flower, to create a shade environment suitable for the growth of the konjac. Intercropping
crops should preferably include long-stalk species to avoid competing for photosynthetic
resources, such as light and CO2, thus the planting density of long-stalk crops should also
be appropriate. Suitable inter-row spacing in sparse forests, tree plantations economic
seeking benefits (e.g., rubber or coffee fruit trees) can also be selected for planting the
konjac, but special attention should be paid to the density of such forests to ensure the
necessary light for the konjac growth (Liu Wenlu et al. 2009).
4.1.4.6.3 Surrounding Environment

The environmental conditions such as vegetation, forest, topography, and geomorphology
around the konjac land can also indirectly influence the growth of the konjac. The pieces
of field close to forests and lush vegetation can get good natural shade, a moist and cool
microclimate, and a high concentration of CO2. The temperature in high-altitude areas is
low, thus, shading should be appropriately managed to ensure the necessary light.
4.1.5 Hazards for Konjac Cultivation

4.1.5.1 Diseases
Disease is the most serious among the various hazards for the konjac cultivation; and soft
rot and southern blight are the most harmful diseases for the konjac, especially soft rot has
become the greatest threat to the konjac cultivation. Konjac planters must have an in-depth
understanding of the konjac diseases and use comprehensive cultivation techniques to
prevent diseases and improve the yield.
4.1.5.1.1 Soft Rot

In Sichuan, Yunnan, Guizhou, Hubei, Hunan, Chongqing, Shaanxi, and other provinces in
China, soft rot is common in areas where the konjac production is concentrated; and the
soft rot in Japan is also very common and serious. In contrast, the A. paeoniifolius, widely
cultivated in India, and the A. paeoniifolius and A. dunnii, produced in Yunnan, are the
starch type konjac with a low incidence of soft rot.
4.1.5.1.1.1 Pathogens The pathogenic bacteria causing the soft rot in the konjac have been
identified as the carrot pathogenic varieties of carrot soft rot, such as, Erwinia aroideae,
Erwinia carotovora subsp. Atroseptica, and Erwinia carotovora subsp. carotovora. In the konjac
cultivation, there is a difference between the varieties that these pathogens infect and also
the degree of pathogenicity among the pathogenic bacteria (Huang Junbin et al. 1999).
4.1.5.1.1.2 Suitable Conditions for Infection Studies have shown that soft rot may occur
between 10°C and 40°C, and it is most common between 22°C and 35°C and is rare below
10°C and above 40°C. The optimum propagation temperature for the konjac soft rot patho-
gen is 25°C–35°C. The pathogen can grow at a pH value of 5.3–9.2, but it grows best at neu-
tral pH. The incidence of the konjac soft rot is the highest at a relative humidity between
90% and 95%. At a given temperature, the higher the humidity, the more serious the dis-
ease. Therefore, the epidemic outbreak of soft rot may easily be caused during a period
of heavy rainfall and high temperatures after the konjac planting, causing wounds in the
epidermis or lack of oxygen (Cui Ming & Li Chuan 2009).
4.1.5.1.1.3 Primary Source of Infection Soft rot bacteria are heterotrophic and need sub-
stances containing carbohydrates that are easy to oxidize as food. They degrade pectate
molecules that bind plant cells together, causing the plant structure to eventually fall apart.
The bacteria can live for up to 40 months in soil rich in compost and organic matter, tak-
ing advantage of abundant carbon sources. When the konjac plants are injured by insects’
gnawing and farm operation, the wounds open an intrusion portal for soft rot pathogen.
Since the konjac corm contains more than 80% water and is crisp and tender, it is easy to
be injured and then be infected with bacteria in the soil. Seed corms carrying bacteria are
the most dangerous primary source of infection. It has been reported that seed corms,
compost, soil, diseased plants, weed roots, insects, and other carriers may be the primary
sources of infection by soft rot bacteria. Any part of the konjac plants can be infected with
soft rot bacteria via wounds (Wang Jian et al. 2009).
4.1.5.1.1.4 Infection and Symptoms After invading the plant, the soft rot bacteria usually
reproduce in the parenchymatous tissue. They first absorb nutrients from the wound part
or intercellular space, and then they destroy cell walls and middle lamella, making the
intracellular water and other inclusions flow into the intercellular space, thus causing his-
tolysis, collapse, and lodging of the plant, until the whole plant shows distinctive signs of
soft rot. Therefore, the seed konjac should be strictly controlled during the konjac planting
to avoid injury or infection of the corm seeds. If the konjac corm is injured and infected
during harvesting, the bacteria may hide in corm tissues and carry over soft rot during
storage and dormancy periods, causing corm rot (He Feng et al. 2011).
4.1.5.1.1.5 Transmission The pathogen overwinters in the soil, diseased plant remains,
and weed roots. During the konjac growing season, it gets attached to the leaflet or the pet-
iole through the soil in the course of farm operations, wind, or rain, and then it invades the
plant to cause disease. When the aboveground part becomes diseased, the diseased plants
become the centre of the secondary source of infection; and the pathogen is transmitted to
adjacent plants via the wind and rain. Meanwhile, the bacteria carried by the underground
corm are also transmitted to adjacent injured corms or plants through the soil.
4.1.5.1.2 Southern Blight

Southern blight is the second serious disease in the konjac cultivation. Although its patho-
gen is different from that of bacterial soft rot, its primary sources of infection and trans-
mission routes are similar, and the prevention and control are also difficult.
4.1.5.1.2.1 Pathogens The konjac southern blight is a fungal disease. The pathogenic
fungus is Sclerotium rolfsii Sacc that endangers the konjac from generation to generation.
The hypha is first white, then turns yellowish-brown, always forms rhizomorphs (special
morphological adaptation root-like structures), and can even tightly pack into a sclero-
tium. The sclerotium is spherical, first white, then dark brown, with a diameter of 1–2 mm
or larger. The sclerotium is highly adaptable to environmental changes and can survive
10 years indoors and 5–6 years in the field. Hyphae rarely produce basidiosomes under
natural circumstances.
The growth temperature is 13°C–38°C; the optimum temperature is 30°C–33°C; and the
optimal pH value is 5.9. If the pH value reaches 8.0, the hyphae cannot grow and the scle-
rotium cannot germinate. The bacteria are highly saprophytic (Ma Qiong et al. 2006).
4.1.5.1.2.2 Symptoms It often invades the plant from the petiole close to the ground by the
hyphae and causes disease. Light pink moist spots are generated at the part invaded by the
hyphae, and then expand around the petiole; later, the tips of the leaflet on the same side of
such spots begin to turn yellow; and the soft rot side of the part of the petiole close to the
ground lodges. White silky hyphae are formed at the part close to the ground, the nearby
ground surface, and the shallow soil layer where the disease develops rapidly, and spreads
to surrounding plants or the ground. Such hyphae get together to form a white sclerotium
with the part close to the ground of the petiole as the centre; and the sclerotium then turns
yellow and dark brown. When the ground surface is dry, since the sclerotium in the deep
soil layer germinates and invades, colour fading and the withering of leaves are observed
before seeing the hyphae or sclerotium. If the petiole is pulled out, hyphae reproduction
can be seen at its base. The disease spots expanding along the petiole invade the corm via
the petiole base and cause rot of the corm or offset tubers. The plants with the early onset
of disease have poor expansion of the corm. If the disease is serious, the yield is signifi-
cantly reduced due to the invasion of the corm. The infected corms are of poor quality, and
thus, can never be used as konjac seed (Yan Kai et al. 2015).
4.1.5.1.2.3 Infection and Transmission The sclerotium and hyphae of southern blight bacte-
ria in the konjac survive the winter in soil, on the corm, in diseased plants and compost, on
fallen leaves and weeds, and in crop rhizosphere. These parts become primary sources of
infection. Most of the sclerotium exists in the 3–7 cm topsoil layer. The sclerotium can also
be found in compost. The southern blight bacteria are saprophytic in the soil, and the new
hyphae germinated from the sclerotium in the following spring can directly invade the
host, generally via a wound. Hyphae can be transmitted through the soil and cause infec-
tion, can be transmitted to healthy plants by contact with the diseased plants, or transmit-
ted over a long distance by bacteria carried by the konjac seed. The bacterium is a soil-borne
disease with a wide range of hosts in nearly 100 families and 500 species of plants, but
Cucurbitaceae, Cruciferae, Leguminosae, and grasses are generally not infected. The dis-
ease is more likely to occur in continuously cropped land or the land with a previous crop
that is a host crop. The disease is serious in the case of unsterilized corms, low-lying land,
acidic soil, loose sandy soil, too much nitrogen fertilizer, and high air humidity, otherwise
the disease is minor (Cui Ming et al. 1999).
4.1.5.2 Insect Attacks on Konjac

The larva of insects for the konjac mainly are aphids, Sphingidae: Theretra oldenlandiae,
Agrius convolvuli, and Clanis bilineata, Scarabaeidae: Anomala corpulenta Motschulsky,
Cicadellidae: leafhoppers, and Gryllotalpidae: mole cricket, mites, and other insects.
The harm of insects to the konjac mainly includes three aspects. First, direct plant damage
is caused by gnawing konjac leaves; second, the corm damage is caused by gnawing the
underground corm of the konjac; and third, the infection, withering, rot, or corm rot of
plants is caused by the direct invasion of the konjac soft rot and other pathogens from the
wounds generated by insects gnawing leaves or underground corms.
4.1.5.3 Meteorological Hazards

4.1.5.3.1 Low and High Temperature Hazards
Although the konjac is a crop that likes a warm environment, too high temperatures may
lead to wilting and desiccation. As for low temperatures, they can easily cause cold dam-
age or freeze injury. This phenomenon is mainly reflected in the frost damage to the seed
konjac, especially in winter or early spring during seed corms storage and transportation.
Moreover, the low temperatures are affecting the planting time, the emergence of the leaves
from the ground, and their expansion. In years with low temperatures and much rainfall,
even if the plant can expand the leaf normally, it may subsequently still easily be infected
with soil-borne diseases. If the seed corms are then dug out, it can be seen that they are rot-
ten. High temperature damages mainly occur in July and August; minor damages cause
physical discomfort; while serious damages cause sunscald of leaves. Particularly high
temperatures and strong sunlight cause a temperature rise of leaves; and the leaves turn
white green with a subsequent browning and withering. High temperatures cause the
accumulated temperature requirements of the konjac to be met quickly, causing the early
ending of growth cycle, lodging, and severe drop in yield. In addition, high temperatures
are often accompanied with a high humidity that can easily cause the outbreak of such
destructive diseases as the konjac soft rot.
4.1.5.3.2 Strong Sunlight

The konjac cannot withstand strong sunlight. Its light saturation point is only around
20,000 lx and the light compensation point only about 2,000 lx. It is easy to exceed its light
saturation point for the monoculture on open flat areas, and this can cause physical dis-
comfort when the exposure to sun and temperature rise cause sunscald of leaves, wither-
ing, and death.
4.1.5.4 Drought Damage

Since the root system of the konjac is shallow, the absorption area is small; and the kon-
jac is not drought-resistant. In recent years, extreme weather conditions are more fre-
quent, especially during the rapid growth period of the konjac corm, i.e., in late July to
August. Droughts occurred in different years in the major growing areas of the konjac
in China, namely, the southern part of the Shaanxi Province, the mountainous regions of
the Sichuan Basin, the western and north-western parts of the Hubei Province, and the
Yunnan-Guizhou Plateau. The konjac is often planted in mountainous areas where irri-
gation facilities are relatively scarce, causing a yield reduction due to drought. Since the
konjac needs less water than other crops, good irrigation effects can be obtained by just
giving a small amount of water at a time.
4.1.5.5 Waterlogging Damage

There are often heavy storms in the wet season, causing topsoil loss or severe soil erosion,
making the konjac seed or its roots exposed or even washing away the konjac seed. Leaves
are seriously damaged by storms and subsequently fade and wither. In the case of continu-
ous rainfall, water accumulates in the soil over time; the oxygen content decreases; and the
respiration of the root system of the konjac is seriously disturbed or even obstructed. As
a result, the roots can easily rot, especially during the high temperatures in summer. Too
wet soil can cause the outbreak of soft rot and other diseases.
4.1.5.6 Damages Caused by Wind and Hail

The konjac grows one leaf in a growth cycle; the leaf is connected by a separation layer at
the top of the corm, but the connection is not firm. In the case of strong wind, the plant is
twisted; the petiole gets tilted; and the leaf backside faces upward. In the case of a violent
typhoon during the corm expansion in July–August, the petiole may be bent to the ground
or broken, causing a severe drop in the yield or even total crop failure that year. If damages
are caused by wind and hail after September, the loss is reduced as the rapid expansion
period of the corm is over. Wind damages are a serious threat for the konjac plantations in
Japan. In the case of hail in the period of emergence of the konjac, the bud is damaged. As
a result, the leaf and the bud cannot expand fully and the yield and quality drop seriously.
If the konjac is damaged after leaf expansion and has large wounds, there is a risk of infec-
tion by soft rot or other soil-borne diseases.
4.1.6 Cultivation Techniques

The objectives of the konjac cultivation are high yield and high quality, and these can
be achieved by following the biological characteristics of the konjac and trying to meet
the requirements of the environment in which it is growing to ensure that the plant is in
sound physiological state and grows vigorously. In addition, planters must also be fully
aware that the biggest threats to the konjac cultivation are still diseases.
There are a number of difficulties in disease prevention and in achieving a high yield
of the konjac. First, no genotype resistant to soft rot and southern blight has been found
among the genetic resources of the KGM type konjac so far. Second, since the major konjac
diseases, such as soft rot and southern blight, have an extensive range of host plants and
are transmitted mainly by seed corms and soil, and the reproduction coefficient of the
konjac is low, it is difficult to sterilize the seed konjac in production, and the effects of seed
soaking in a fungicide solution and similar methods are not sufficient. Third, the envi-
ronmental conditions for the spread of major diseases are mostly matching the suitable
conditions for the growth of the konjac plant. This coincidence is difficult to manage and
solve. However, since the major diseases of the konjac, such as soft rot and southern blight,
are transmitted by seed corms and soil, there is a scope for the elaboration of the appro-
priate prevention and control measures (Hayashi 1988). Production practices have proved
that diseases can be controlled by the implementation of measures such as planting in the
most suitable areas, adopting crop rotation, breeding disease-free seed corms, steriliza-
tion of the seed konjac and soil, intercropping, shade, soil surface coverage, ridge cultiva-
tion, timely pest control, timely harvesting, and protective storage. In contrast, continuous
cropping, use of seed corms infested with pathogens, high temperatures and rainfall, poor
drainage, excessive nitrogen fertilizer, and insect damage in the cultivation area can easily
promote disease development (Liu Peiying & Zhang Shenglin 2000; Chen Dunqiao & Yang
Qinglin 2000).
4.1.6.1 Selection of Areas and Plots for Cultivation

4.1.6.1.1 Selection of Areas for Cultivation
There is a large zone suitable for the konjac to the south of the Qinling Mountains (a major
east-west mountain range in southern Shaanxi Province) in China. At present, only a small
part has been developed and utilized. Therefore, the konjac industry should select cultiva-
tion plots in the particularly suitable areas, in particular, in those included in the ‘Regional
Planning for Planting Konjac in China’. The blind development in unsuitable areas is dis-
couraged. Since the konjac has strict requirements for environmental conditions and the
environment cannot be artificially changed, cultivation on inappropriate sites requires
more efforts, produces less gain, and may end with failure.
According to the ‘Regional Planning for Planting of Konjac in China’, the lower valley
of the Yarlung Zangbo River, the tropical district in southern Yunnan, and the Limuling
Mountains in Hainan have a subtropical humid climate and are particularly suitable area
for planting the konjac; the Yunnan-Guizhou Plateau (including the southern Sichuan
Plateau), mountainous areas around the Sichuan Basin, Nanling Mountains (including
Western Hubei and western Hunan parts of the Wuling Mountains, Luoxiao Mountains,
and Wuyi Mountains), and the mountainous areas to the south of the Nanling Mountains
are warm and humid and are the most suitable for planting the konjac; the Qin-Ba
Mountain areas to the north of the main ridge of the Daba Mountain area, and to the west
and northwest of the Hubei Province are suitable for planting the konjac.
When selecting specific planting bases, the konjac should be planted in areas where
the mean monthly temperature in May–October is higher than 15°C, the mean maximum
temperature in July–August does not exceed 32°C, the total rainfall is above 200 mm, the
relative humidity is between 80% and 85%, and there is natural or artificial shading. In the
growing period of the konjac in May–October, areas with a longer period of a daily tem-
perature of around 25°C are more conducive to the growth and development of the konjac,
the yield is higher and the risk of disease is small (Zhang Zhongliang et al. 1998).
Since the konjac has strict requirements for environmental conditions, its suitable areas
are obviously and comprehensively related to the altitude. Except for a few low-altitude
areas with an adjustment of the ecological and environmental conditions by forest and
water, most of the suitable areas for the konjac in China are in the higher-altitude moun-
tain or hilly areas. The altitude should be selected according to the latitude, mountains,
rivers, topography, landform, vegetation, and climate in the area, taking advantage of the
three-dimensional climate in 500–2,500 m mountainous areas. According to years of prac-
tical experience, the upper altitude limit for rice cultivation in the area where the konjac is
to be planted generally is the lower altitude limit for the growth of the konjac. For instance,
the suitable altitude of the south foothill in the Qin-Ba Mountain areas is about 1,000 m,
but the konjac should be developed at an altitude of about 800 m on the north foothill; the
areas in the Yunnan-Guizhou Plateau with an altitude of about 2,000 m are suitable for the
konjac development. In areas where the environmental conditions of the konjac, especially
temperature, can be met, the altitude should be properly selected. For example, the konjac
can be cultivated at an altitude of nearly 1,000 m in the Jinsha River Basin, mountain areas
around the Sichuan Basin, and the northwest Hubei mountains, but diseases are signifi-
cantly reduced by selecting the altitude of 1,200–1,500 m.
4.1.6.1.2 Selection of Environment for Cultivation Plots

In areas with appropriate overall environmental conditions, the specific environment of
the plot should also be selected. The cultivation plots should be cool and moist in the sum-
mer to facilitate the growth of the konjac, and warm and dry at the end of the growing
season to help the corm harvesting and storage. Therefore, the plots with a water source,
irrigation conditions and good drainage, and without severe soil erosion caused by rain-
storms in the summer in sloping and leeward areas blocked by a chain of mountains or
shaded with trees and with semi-shade conditions should be selected. The selection of
a shady slope in mountainous areas should be considered in combination with the local
temperature: if the temperature is close to the optimal temperature for the growth of the
konjac (25°C) during the growing period, and there is no damage caused by high tempera-
tures above 35°C, the plots on sunny slopes should be selected to obtain sufficient sunlight
and increase the yield; and if there are high temperature threats (above 35°C), the plots on
shady slopes with a relatively small amount of sunlight should be selected.
4.1.6.1.3 Soil
For the optimum growth of the konjac, a fertile soil with a top layer of more than 30 cm,
rich in organic matter, well drained, but with good water and nutrients retention, and
neutral pH should be selected. Loam or sandy clay loam is preferred. A sandy soil with
poor water retention and too heavy clay soil with poor drainage and aeration are unsuit-
able for planting the konjac. The soil with inadequate organic matter and lacking granular
structure, having insufficient water retention, poor drainage, and inadequate nutrient con-
tent and ventilation should be avoided. In such soil, the konjac plants do not grow vigor-
ously and may easily be infected with diseases. On the basis of the selection of soil, the
appropriate amount of fertilizer should be applied to improve the soil fertility. The way of
improving the uptake of fertilizer and improving the soil structure is through the applica-
tion of soil amendments (improving the permeability and water retention characteristics
of the soil) and tillage. In addition, a large amount of high-quality thoroughly decomposed
and pathogen-free compost may be applied. The ideal soil for planting the konjac has strict
requirements with regard to physical and chemical characteristics (GPAIA 1991), as shown
in Table 4.1.
4.1.6.2 Cultivation Pattern and Cropping System

4.1.6.2.1 Cultivation Pattern
4.1.6.2.1.1 Natural Growth Method The konjac is a perennial plant originating from the
undergrowth of forests in tropical, rainy climates and completes its life cycle in 3–4 years.
Subsequently, its natural population is a mix of 1- to 4-year-old plants. Since the occurrence
and outbreak of the konjac soft rot and southern blight are closely related to plant injuries
caused by farm operations, the plants living in their natural environment in the wild have
a lower risk of injury and infection with the pathogens. Natural growth method (more on
that in Section 4.2) is adopted in Japan, China, and some countries in Southeast Asia.
TABLE 4.1
Physical and Chemical Characteristics of Soil Suitable for Konjac Cultivation
Characteristic
Type Characteristic Value Remarks
Physical Topsoil depth >20 cm

characteristics Density Plough layer 15 mm
Depth below plough 20 mm
layer
Underground water level >60 cm
Porosity Solid phase volume 40%–60%
Liquid phase volume 20%–30%
Air phase volume 20%–30%
pH value 6.0–6.5 Water measure
Chemical Conductivity 0.3 ms/cm
characteristics Replacement CaO 150–400 mg Measured with 100 g air-dried soil sample
Replacement MgO 25–60 mg Measured with 100 g air-dried soil sample
Replacement K2O 150–402 mg Measured with 100 g air-dried soil sample
Available P2O5 150–403 mg Measured with 100 g air-dried soil sample
CaO/K2O 3–8
MgO/K2O 1–2
CaO/MgO 4–6
The natural growth method can be adopted in areas that are the most suitable for the
growth of the konjac, have an annual mean temperature of 13°C–18°C without intense heat
and intense sunshine in summer and without frost in winter, with a thick fertile topsoil
layer, high organic matter content, and shading provided by deciduous trees. Generally,
no tillage is required. Only an appropriate amount of organic fertilizer is supplemented to
ensure the high quality of the corm; corms of 3-year-old plants are harvested according to
the thickness of the petiole, with 1- to 2-year-old corm left in the soil for further growth.
Selected harvesting is conducted in successive years without tillage.
4.1.6.2.1.2 Field Cultivation Corms, lateral corms, and offset tubers of the konjac are
planted in the field according to the type of planting material and age. After harvesting
all corms, the 2- to 3-year-old corms are sold for processing. The 1- to 2-year-old small
corms, offset tubers, and lateral corms are used as seed material after storage in winter
for further cultivation. The injuries of corms may be caused during drying, storage, and
transportation of seed konjac and inadequate field management. The disease incidence is
much higher than when practicing the natural growth method. Moreover, the decrease
of corm quality and yield may be caused by improper application of pesticides and fertil-
izers. Between the two cultivation patterns, the natural growth method is very consistent
with the biological characteristics of the konjac and produces a high yield and quality.
However, with the development of production and the expansion of the cultivation area,
the production pattern has gradually turned to field cultivation. After years of experience
in field cultivation practice, relatively efficient cultivation methods have been developed.
For example, some areas in the Yunnan Province, China, adopt 2-year natural cultivation.
This means that in a suitable natural environment that is suitable for the growth of the
konjac, small seed corms are planted in the first year. After appropriate fertilization, cul-
tivation in deep furrows and high mounds is carried out without harvesting in autumn.
In the winter, the plants are properly covered with soil or with green manure to protect
them from frost. In the spring, after applying fertilizer, the dry green manure is laid on
the plant bed. The harvesting is conducted in autumn including large size corms for pro-
cessing, commercial seed corms (the second-generation seed), and lateral corms (the first-
generation seed). In Langao County (Ankang, Shaanxi Province) and other regions, the
konjac is planted under sparse woodland that has been converted from farmland to forest
to restore the conditions similar to the natural growth method (Zhang Yang 2017).
4.1.6.2.2 Cropping System

4.1.6.2.2.1 Crop Rotation Crop rotation is of great significance to the konjac cultivation.
The debris of the konjac that are left in the soil may contain the bacteria of the main dis-
eases for the konjac, such as soft rot, southern blight, root rot, and bacterial leaf blight and
become the primary source of infection. Crop rotation is the most fundamental way to
block the transmission of pathogens. Generally, each rotation is at least 3 years. The plot
considered for crop rotation should avoid a previous crop of Cruciferae, Solanaceae, gin-
ger, and other tuber crops as much as possible. The planting of gramineous and legumi-
nous crops is relatively safe. Dryland cultivation including an appropriate crop may be
implemented within rotation in regions where conditions permit.
4.1.6.2.3 Intercropping System

1. Principle of intercropping: When the konjac is intercropped with trees or other crops,
the leaf area index will be increased and the corm yield per unit area improved.
Moreover, as the konjac should avoid strong sunlight, intercropping will provide
the konjac with more shade. Thus, the risk of high temperatures caused by strong
sunlight will be reduced. The key to successful intercropping is to control the
degree of shading. With moderate plant density and shade, the konjac and the
other crops can both benefit. When applying appropriate intercropping, the corm
yield per unit area is much higher than in monoculture. The shade density of
40%–60% is appropriate and can both inhibit diseases and stimulate photosynthe-
sis, resulting in a higher yield.
2. Crop selection and distribution of intercropping: The shade plant selected for inter-
cropping should be higher than the konjac. The height of 2- to 3-year-old A. konjac
generally is 1 m, and that of the A. albus is 60 cm. Therefore, the best intercrop-
ping crops are young deciduous trees and long-stalk crops. Among suitable trees
are: Eucommia ulmoides, Paulownia sp., deciduous fruit trees, mulberry, Camellia sp.,
and Ricinus communis. The examples of crops are: sorghum, corn, amaranth, jute,
and sunflower. Thus, trees and long-stalk crops can obtain more sunlight as an
upper layer and the konjac can be properly shaded as a lower layer. The konjac
generally emerges in mid-to-late May and gets mature and decays in late October
or early November. After September, the temperature and illumination gradually
decrease, thus, the shade should be decreased by removing the lower leaves of the
sorghum, corn, sunflower, and other crops at the later stage of growth to increase
the sunlight for the konjac.
4.1.6.3 Reproduction Methods

4.1.6.3.1 Reproduction by Seed
Generally, this method is only used for increasing the amount of seed material and for
breeding of the later generations in crossbreeding. The later generations have greater
variation and, thus, provide more selection opportunities. However, the time when the
commercial konjac is obtained is usually 1–2 years later than the reproduction with off-
set tubers. In recent years, with the rise of the development of the A. muelleri, various
regions have started to breed the seed of the A. muelleri. Such seed has gradually been
produced and used for planting in Yunnan Province, China, Thailand, Laos, Myanmar,
and Indonesia.
4.1.6.3.2 Reproduction by Offset Tubers

Offset tubers are also known as rhizomatous offsets and are good reproduction mate-
rial in large quantity. Different varieties develop a different number of offset tubers on
each corm from three to more than ten offset tubers (such as A. albus). Corms of different
ages also have a different number of offset tubers. Generally, 1-year-old corms have fewer
offset tubers, while 2- to 3-year-old corms have more offset tubers. When the offset tubers
of the A. konjac get mature, the end connecting it to the corm usually shrinks gradually,
forming the ‘umbilicus mark’ of offset tubers on the corm. Then the offset tuber breaks
away from the corm and becomes an independent, small offset tuber. Even after reaching
maturity, the offset tubers of A. albus do not break away from the corm and this needs to
be done manually. Moreover, since the offset tubers of the A. albus can be as long as 20 cm
and each segment has buds, such segment can be used for reproduction.
4.1.6.3.3 Reproduction by Whole Corm and Cuttings

Generally, 1-year-old corms are planted for reproduction as whole corms; 2-year-old corms
(below 500 g) can also be planted as whole corms. The corms above 500 g can be cut for
reproduction to increase the reproduction rate and are usually cut with a thin knife from
the terminal bud downwards into 4 ~ 6 pieces, each weighing about 100 g. The terminal
bud must be cut to destroy its apical dominance and promote the lateral leaf buds on the
cuttings to germinate as leaves. Attention should be paid to the air drying and healing of
the cutting wound to prevent infection; an appropriate temperature and humidity should
be maintained during the storage of cuts. If corms are cut right before spring planting, the
budding period will be delayed, affecting the yield. Therefore, corms should be cut before
storage.
4.1.6.3.4 Reproduction by Tissue Culture

The tissue culture technique is used for in vitro asexual reproduction to accelerate the
reproduction and, thus, the problem of a low reproduction rate can be fundamentally
solved. The corms, offset tubers, petioles, internal buds (on the top of the corm), flower
stalks, or stem tips of the konjac can form a callus, but the effects of the tissue culture and
rapid reproduction with corm cuts as explants are the best. If stem tips are used as explants,
a sterilization treatment can be conducted at the same time. The callus induction rate of
the A. konjac and A. albus is relatively high. When the bud begins to grow a scaly leaf, the
seedling that has grown root is transplanted to a seedbed rich in humus. The seedbed is
covered with plastic film to preserve moisture. The success rate of transplanting can reach
100% if properly managed (Zhang Xingguo et al. 1993). Small corms weighing between 3
and 6 g can be formed at the end of the growth cycle. At present, there also are reports on
altering the tendency of the callus to form tissue culture seedlings towards directly form-
ing small corms by means of temperature changes and induction of hormone ratio (Dai
Qingtang et al. 2011).
4.1.6.4 Seed Konjac Selection and Treatment Before Planting

4.1.6.4.1 Seed Konjac Selection
The quality of seed material is the key factor affecting the potential severity of disease, as
most diseases are transmitted via seed konjac. Moreover, the quality of seed material has
great impacts on the early growth of leaves and roots. The sources of seed konjac should be
the plots without diseases. Injuries, infection by diseases, and other undesirable conditions
of seed konjac should be checked during harvesting and before drying and warehousing.
The seed konjac with minor cuts that heal after air drying can still be used. When small
corms and offset tubers are used as seed konjac, small corms and offset tubers should be
strictly examined, and any damaged, crushed, or dried seed konjac or seed konjac with
frost damage, mildew, or diseases and pests should be removed to ensure the quality of
seed material. In terms of the weight of the seed konjac, the seed material of the A. konjac
should be less than 500 g and that of the A. albus below 100 g. The seed konjac should be
segregated and planted according to the variety, age, and weight.
4.1.6.4.2 Sterilization of Seed Konjac

4.1.6.4.2.1 Sterilization by Drying During the sorting of the seed konjac, the seed material
can be dried by the sunlight on sunny days until the epidermis is slightly dehydrated; dur-
ing this process, the seed konjac is sterilized by the sunlight.
4.1.6.4.2.2 Sterilization by Anti-Microbial Agents Such agents as streptomycin (wettable

powder), formalin, copper sulphate, potassium permanganate, carbendazim, and a few
other anti-bacterial compounds are generally used for sterilization of seed material.
They are mainly applied by spraying or soaking. The abovementioned agents are usually
applied separately and diluted according to the instructions of the manufacturers. If soak-
ing sterilization is adopted, the soaking time should be 20–30 minutes. The seed konjac
usually needs to be dried after spraying and soaking sterilization to ensure that the agent
is fully attached on the surface of the seed material.
4.1.6.5 Plot Preparations

4.1.6.5.1 Soil Preparation
The deep ploughing of the field with a layer depth of 25–30 cm should be conducted before
the winter to ensure that the topsoil is fully mixed. In a few areas, the rain and snow
in the winter can be used to kill some bacteria and pests. Before the planting begins in the
spring, weeds should be removed; and 1,000 kg thoroughly decomposed farm manure or
organic fertilizer and 25–30 kg potassium fertilizer are applied to each mu (1 ha = 15 mu).
Then, 2.5 kg sulphur powder mixed with 100 kg plant ash, or 100 kg quick lime, is evenly
scattered on each mu for soil disinfection. Finally, fine tillage and raking of the soil level
are carried out (Yang Hongmin et al. 1989).
4.1.6.5.2 Ditching and Ridging

Ditching and ridging are conducted with 0.7–1.2 m spacing between rows. The root sys-
tem of the konjac is shallow and requires adequate oxygen, thus, good drainage must be
ensured; and in areas with plenty of rain, the konjac is planted with deep ditches and a
row spacing of about 1 m and a ridge height of 15–20 cm. In areas with less rain in the
summer and autumn, the konjac can be planted with wider spacing and shallow ditches.
4.1.6.6 Planting
4.1.6.6.1 Planting Time
The konjac is generally planted after the termination of the physiological dormancy period
of the corm and when the mean temperature rises to 12°C–14°C and the minimum tem-
perature is about 10°C. The konjac is generally planted in mid-to-late March in a warmer
climate, in April, in areas at higher altitude and northern latitude in China, and in early
May, in northern areas in Japan. The conventional planting period in Korea is around April
20; and the yield can be increased if the planting is advanced to April 5 and covered with
mulch (Lee 1992). In low mountainous areas where there is a warm winter without frost in
southern China, seeding planting in winter can be carried out immediately after the har-
vest in November–December; big corms are harvested and small ones left; and small corms
and offset tubers are evenly buried in the soil for natural germination in the next spring.
4.1.6.6.2 Planting Density

The plant spacing within a row is usually such that the leaves overlap about one third after
the leaf expansion of the konjac plants. Plant density is closely correlated with the size of
seed material, the larger the seed konjac, the larger the planting spacing. In order to ensure
ventilation and make it convenient for field management, wide spacing between rows and
narrow spacing within a row are generally adopted. Seedlings are usually planted in
biodegradable nursery pots, with 2–3 plants/pot. Lateral corms are usually planted with
12–15 cm spacing within a row. The spacing between rows for the second-generation seed
konjac is usually six times the diameter of the corm and the spacing within rows four times
the diameter (Watanabe Hiroshi 1979).
4.1.6.7 Field Management

4.1.6.7.1 Weeding
The root system of the konjac is shallow and spreads horizontally in the upper layer of
the soil. Thus, intertillage (tillage or cultivation between plants) aiming at weeding may
easily cause root damage and diseases. In weedy plots, weeds should be destroyed with a
pre-emergent weed killer (approved by the phytosanitary authority) or removed manually
after planting in the spring and before emergence. After the konjac comes up out of the
ground, weeds should be removed manually without damaging any part of the plants.
4.1.6.7.2 Soil Loosening

The konjac begins to germinate and emerge from the ground about one month after plant-
ing. This is the appropriate time to carry out shallow ploughing (5–10 cm deep) to break
up the crust located on the surface of the soil to increase the soil permeability in order to
improve the gas and moisture transfer. The timing is important, as this operation has to
take place before the root system is fully developed in the top layer of the soil. Soil loosen-
ing and fertilizing can usually be implemented at the same time.
4.1.6.7.3 Soil Surface Coverage

Soil surface coverage can prevent rainstorms from directly washing the soil, causing fertil-
izer loss and root exposure. It protects the granular structure in the soil, adjusts soil tem-
perature and humidity, prevents weeds from expanding rapidly, reduces risk of diseases,
and improves the yield. Pine needles, fallen leaves, rice straw, dried bamboo leaves, and
corn stalks are usually used as mulching materials in south China. In Japan, oat or barley
are planted on the side of the konjac furrow and straw is cut to cover the soil surface with
a covering thickness of 4–6 cm.
4.1.6.7.4 Fertilizing
In China, the first application of fertilizer after emergence is usually in early June to pro-
mote the growth of the aboveground parts of the konjac. The second application is in
early July to promote the growth of the underground parts. The corm begins to expand
quickly in mid-to-late July and absorbs a large amount of nutrients in August–September,
but the extensive application of nitrogen fertilizer in the middle and later periods causes
the excessive growth of the aboveground part and is unfavourable to the accumulation of
active components in the corm.
4.1.6.7.5 Pest Control

In addition to the use of disease preventing agents during soil and seed konjac steriliza-
tion, pest control should be continued during the growth of plants according to the devel-
opment of diseases and pest conditions.
4.1.6.7.5.1 Physical Methods Diseases can be prevented and controlled by grading of the
seed konjac, crop rotation, intercropping, timely removal and burying the affected plants,
and other cultural disease control techniques. Infested plants should be removed manu-
ally as soon as possible. Also, steam treatment of the top layer and deep ploughing in the
winter are other ways of physical control of diseases and pests.
4.1.6.7.5.2 Chemical Control The results of the tests on the effectiveness of commonly used
agents on southern blight and soft rot showed that thiophanate methyl, mancozeb, and
metalaxyl provide the best control of southern blight, while carbendazim, anti-toxic alum,
and streptomycin provide the best control of soft rot (Cui Ming et al. 2001). The Bordeaux
mixture is generally used for disease prevention in Japan. Planned spraying is carried out
monthly from the early stage of leaf expansion; which is especially effective in prevent-
ing disease in leaves that tends to develop disease after a typhoon. In recent years, there
have been reports of successful control of the konjac diseases by biological pesticides (Cui
Pengfei 2016).
The larva of sweet potato noctuid moths, Sphingidae, and Clanis bilineata may be cap-
tured manually and may be sprayed with a diluted phoxim oil concentrate or decis, a
broad spectrum pyrethroid insecticide, when the pest attack is serious. In terms of the
methods to control grub or mole cricket, in addition to avoiding the use of undecayed
manure or compost, the thoroughly decomposed organic manure and light may be used
to kill adults. If larvae damage is found, a diluted phoxim oil concentrate may be used for
root treatment.
4.1.6.8 Harvesting
In China, after the end of September, the corm gradually enters the dormancy period,
leaves tend to stop growing and gradually turn yellow and reach the senescence stage.
The maturity and senescence stage of leaves takes about 10 days and commercial size
corms may be harvested from this time. Harvesting should be conducted on sunny days
and corm injury should be avoided during harvesting.
4.1.6.9 Storage of Planting Material

The growing period and storage period of the seed konjac are about half a year, respec-
tively. Although high-quality seed konjac is produced, the quality and shelf life of the seed
konjac may also be reduced due to improper storage and management. As a result, the
growth and yield in the following year will be affected.
4.1.6.9.1 Principle of Konjac Corm Storage

4.1.6.9.1.1 Physiological Changes During Storage The physiological changes of the konjac
corm during storage after harvest can be divided into four stages. The first stage is the
after-ripening period, i.e., the early storage period. It is the early dormancy period of the
corm. In China, this period is usually from the maturity and senescence of the konjac leaves
to early December. Following the after-ripening, the corm epidermis is less permeable; the
evaporation intensity and respiration gradually weaken to turn into the dormancy state.
The second stage is the deep dormancy period, the activity of various enzymes and the
respiration in the corm weaken. In such cases, the corm is not sensitive to temperatures and
does not germinate even under favourable environmental conditions. The third stage is the
dormancy release period generally from late February to March. At this stage, the activity
of catalase gradually decreases, and that of polyphenol oxidase gradually increases; and the
respiration is weak. The bud can germinate if the ambient temperature reaches the thresh-
old temperature of the konjac germination. The fourth stage is the bud elongation period.
After March, the dormancy of the corm is terminated; and the corm can germinate to form
a plant under suitable environmental conditions (Sun Yuanming et al. 1995).
4.1.6.9.1.2 Curing Before Storage The effects of the konjac storage are closely correlated to
the temperature, humidity, ventilation status, and other environmental conditions during
the storage, with proper field management, timely harvesting, and proper pre-treatment
before the storage as the preconditions for safe storage. The injury of seed konjac is the
most important reason for the rot during storage and after planting; therefore, special
attention should be paid to avoiding the injury of seed konjac from harvesting. The corm
that is just harvested, especially that is harvested prematurely and is large in size, has high
water content, tender epidermis, and crisp meat, and may easily have external wounds and
internal cracks, causing the invasion of soft rot bacteria and rot. Therefore, curing of the
corms should be carried out (Holcroft 2018). The objective of curing is to remove the water
on the surface of the corm to obtain the epidermis suberization and to heal the wound.
Curing begins with sun drying in the field for a day on sunny days after harvesting. After
that, all dirt should be removed and corms spread out in a ventilated place sheltered from
the rain and air dried at 15°C–20°C. The degree and duration of air drying are determined
by the degree of weight loss of the corm. Generally, a 15% loss in weight is required. Two-
year-old corm should be dried for 15–20 days. For offset tubers, the curing objective can be
achieved after 5–10 days of air drying.
4.1.6.9.1.3 Conditions for Safe Storage In order to prevent the formation of ‘aged buds’ and
the occurrence of freeze injury, the storage temperature must be maintained at 8°C–10°C.
At the late stage of seed konjac storage, the temperature may vary between 12°C and 20°C
to promote the germination of the terminal bud. The relative air humidity suitable for
the konjac storage is 80% and should not exceed 90% or fall below 60%. If the relative air
humidity is below 60% for a long time, corms may shrivel easily. The high range of rela-
tive humidity of the air should be maintained in order to reduce the natural loss of corm
moisture, but condensation should be avoided in order to avoid the risk of rot that would
cause considerable loss. This risk is considerable if storage occurs at a high temperature
(30°C–32°C) and high humidity (above 90%), when soft rot spreads easily. Since the corm
still continues to metabolize by breaking down nutrients and releasing heat and water in
the storage period, the temperature and humidity of the storage environment increases,
but the excessive increase in temperature and humidity is unfavourable to the storage.
The simplest method to adjust the temperature and humidity is ventilation. Since the
outdoor temperature is low, ventilation can decrease the temperature and humidity (Liu
Peiying & Wang Yulan 1990).
4.1.6.9.2 Methods of Storage

4.1.6.9.2.1 Storage in Insulated Warehouse A ventilated insulated warehouse should be
built for the large-scale storage of the seed konjac for elite varieties. Stacking racks for
8–10 layers need to be provided in the warehouse. The spacing between layers should be
about 20 cm. The lowest layer is just above the ground. Metal strips or wooden or bamboo
battens are nailed on racks for storing seed konjac, leaving some space for ventilation.
In regions where the temperature falls below 0°C in the winter, a furnace or electric heater
apparatus and ventilator may be installed in the warehouse. A humidifying system may
be installed to adjust the relative humidity, if required.
4.1.6.9.2.2 Simple Indoor Tempering Storage A layer of dry river sand is spread on the floor,
then a layer of seed corms is placed. The corms are again covered with dry river sand and
3–4 layers are formed. The top layer and the four sides are covered with dry straw for
insulation. A layer of dry rice husk, wheat glumes, leaf litter, or straw may be spread on a
ventilated dry floor; then a layer of seed corms is placed; and the two steps are repeated
3–4 times. In a few areas, a cellar may be created on a rocky hillside when seed corms
are placed. They are sterilized with formalin, potassium permanganate, and other agents.
During the dormancy period, the cellar door should be tightly closed, but with vent holes
above the cellar door. Once the temperature rises in the spring, the cellar door may be
opened in the morning for ventilation and closed at night. During storage, the konjac
should be inspected and rotten coms removed promptly.
4.1.6.9.2.3 Outdoor Piled Storage In areas where there is no risk of frost injuries, a dry
place with high terrain, loose soil, good drainage, and exposure to the sun may be selected.
Sorghum stalks are spread on the ground, then a layer of corms is placed, next, loose dry
soil is scattered, and after that, another layer of corms is placed. Such steps are repeated
several times. The top is covered with loose dry soil and sealed with a plastic sheet for pro-
tection against rain. The film should be opened on sunny days for ventilation and closed
on rainy days to prevent the entrance of rainwater. The ventilation should be increased
after arrival of warm days in the spring.
4.1.6.9.2.4 Open Field Protection for Overwintering In regions with frozen ground in the
winter, after the decay of the aboveground parts of the plants on the plots reserved for
seed corms, the holes left after the decay of petioles are filled with soil. Then the soil on the
planting bed surface is covered with mulch (straw, leaves) or a plastic film. In some areas,
leguminous green manure crops can be sown around October on the planting bed of the
konjac in order to use them as mulch for overwintering.
4.2 Agroforestry-Based Production System in Indonesia

4.2.1 The Agroforestry Environment and Its Sustainability
4.2.1.1 Konjac Agroforestry System and Environment
The main konjac species grown commercially in Indonesia is the Amorphophallus muelleri,
called ‘porang’ in Bahasa Indonesian. The konjac had been long known as the produce
from the forest in Indonesia since its processing for commercial food was introduced by the
Japanese people living in Indonesia as early as the 1980s. For a long time, there have been
small- and medium-sized industries processing fresh tubers and dried chips into konjac
noodles for breakfast food or konjac tofu. The konjac agroforestry system has been started by
interweaving it with the area of the state industrial forest, namely, Perum Perhutani a state for-
est holding company mainly producing logs. The konjac is planted as an intercrop under the
shade of wood trees of the Perhutani forest. In the operation level, Perhutani has 57 KPHs or
Kesatuan Pemangkuan Hutan (Forest Management Unit) over all the provinces in Indonesia, but
the konjac is mostly cultivated in KPHs in Central and East Java. The cooperation between the
KPHs and the farmers’ community is further established through the policy called Pengelolaan
Hutan Bersama Masyarakat- PHBM (Community-Based Forest Management).
One of the KPHs located at Saradan, Madiun, East Java called KPH Saradan became the
first centre of the konjac cultivation and production. The area of production in Saradan
reaches presently from 550 to 900 ha at the village of Klangon and from 200 to 300 ha at
the village of Pajaran. Lately, the konjac cultivation has been largely disseminated sur-
rounding the forest into other KPHs at Central and East Java by the interest of the farmers’
community. These KPHs, however, should be distinguished from the recent initiatives of
the government to develop Kesatuan Pengelola Hutan (as well literally translated to Forest
Management Unit and abbreviated KPH), which are aimed to regulate the commercial for-
est managed by the private sector (Hartojo 2018: personal communication).
The predominant trees in the lower land part of the agroforestry are teakwood (Tectonia
grandis Linn. f.), while in the higher land part they are mahogany (Swietenia mahagoni King)
and sono keling (Dalbergia latifolia Roxb.), with the variant of sonobrit and sono sungu. Some
of the trees are 50 years old or older, but others are renewed trees substituting the previous
trees that have already been harvested during logging.
Farmers learned that the konjac plants growing under sono trees provide a better yield
compared to those growing under teakwood and mahogany. In the KPH Saradan, the area
of the forest is divided into a teakwood area, mahogany area, sono area, and mixed trees
area. The size of the forest area under mixed trees is the largest, followed by teakwood,
sono, and mahogany, which is the smallest in size.
4.2.1.2 Agroforestry Sustainability

The goals of KPH Saradan in initiating the agroforestry system with the konjac farming
under the PHBM scheme are as follows:
1. Sustainability of the forest trees which produce high commercial value wood pro-
tected by the government with strict regulations in logging.
2. Sustainability of the forest to maintain the green environment and to avoid green-
house effects.
3. Sustainability of the forest environment with the community dwelling in its sur-
roundings to avoid illegal logging and social conflicts.
4. Sustainability of the prosperity of the community in the forest surroundings.
When the community surrounding the forest becomes konjac farmers, they assist the KPH
in maintaining the forest and the high value wood producing trees because without the
shade of the trees, the konjac growth will fail. At the same time, the community has good
income and is leveraged above the poverty line.
The KPH with the farmers’ community in the villages surrounding the KPH forest
formed an organization called Lembaga Masyarakat Desa Hutan(Forest Village Community
Institution) or Masyarakat Pengelola Sumberdaya Desa Hutan (Forest Village Resources
Management Community). The community institutions elect the leader whose functions
are: (1) coordinating the harvesting time, and formulating and making decisions on the
market price of the produce, which are crucial in the system and (2) liaising between the
community institutions and the KPH. When the system is established, and all parties –
individual farmers and the KPH – reach a win–win solution, both the community institu-
tions and the KPH prefer not to change the leader and feel safe to be coordinated by the
same person for a long time.
This sustainability has been proven in many years of konjac growing under the agro-
forestry system at KPH Saradan, as well as from it to other KPHs at Central and East
Java Provinces. The production area has been increasing from 315 ha at KPH Saradan
in 1998 (Purwadaria 2001) to 1,655.3 ha at Saradan, Jember, Nganjuk, Padangan, Madiun,
Bojonegoro, and Mojokerto in Central and East Java Province (Hidayat et al. 2013). At KPH
Saradan alone, the statistics showed that from 315 ha in 1998, the konjac area has grown to
a 750-ha harvest area in 2017.
4.2.2 Konjac Growth Cycle

4.2.2.1 Cultivation
Cultivation of the konjac plants in Indonesia is managed by the farmers’ community liv-
ing in the forest surrounding the KPH located in East Java and Central Java. The konjac
is planted under the shade of the wood trees and grows ideally under the condition of
estimated 50% sunlight intensity (see Figure 4.1). The altitude of the land is 100–700 m
above the sea level, and the selected land is slightly inclined to allow the surface water
to flow down without flooding the plants. Initially, the farmers obtained the bulbils and
seeds from the konjac plants growing wildly, and, after a while, they could scale up their
cultivation fields bit by bit by using the bulbils and seeds from their own cultivated plants.
Planting could be done in three ways: two vegetative ways by planting the bulbils and
planting the tubers and one generative way by planting the seeds. The planting distance
for bulbils and plantlets originating from seeds is 25 × 25 cm, while for tubers weighing
around 200 g, it is 50 × 50 cm. Large size tubers will be cut into two or three pieces before
planting (Hidayat et al. 2013). Farmers do not have a habit of providing fertilizer for the
konjac plants for unknown reasons. It could be that planting under the wood trees dis-
courages them, thinking that the fertilizer for the konjac plants may not be suitable for
wood trees, and vice versa.
4.2.2.2 Harvesting of Tubers

The harvest is done once a year for the 3-year-old tubers, sorted out by the size of the plant
stems reaching 5 cm diameter or more. Harvesting is done after the plants died and no
stems are seen above the ground. To distinguish the site for digging the tubers, the farmers
put a straw as a mark to find the selected mature ones.
The time for harvest falls between May and August. Plants growing on the lower land
will mature earlier than the plants growing on the higher land. The harvested tubers usu-
ally account for 1/3 to 1/2 of the total number of plants growing on the plot. The rest of the
FIGURE 4.1
A. muelleri growing under trees in KPH Saradan (Central Java).
plants will be remaining in place and will grow again in October and eventually will die
in April, one month before the harvest in the following year.
4.2.2.3 Replanting
To maintain the sustainability of the porang production, farmers need to do replanting to
substitute the plants they harvest. There are three ways of replanting: the most common is
by using the plant bulbils since the plants growing from bulbils will lead to the production
of ready-to-harvest tubers in 2 years, while the other alternatives are by using the plant
tubers, and the plant seeds that commonly need 3 years’ time for reaching the mature
stage that is suitable for harvest. Farmers believe that the rate of plant growth from bulbil
replanting is 100%, while from good seeds is 94%. Replanting by using tubers is avoided
by farmers for the reasons that tubers are the end product for sale, while bulbils are abun-
dant. The other advantage of replanting by bulbils is that the replanting is much simpler
than replanting by seeds.
However, harvest might leave some small tubers accidentally removed from the soil
along with the targeted big ones. Tubers with the size range of 4–10 cm diameter will
be replanted by the farmers. Tubers with 10 cm diameter are commonly divided into
four pieces and then each piece will be planted separately.
4.2.2.3.1 Replanting by Bulbils

Bulbils are collected after dropping on the ground in April, usually after the plant stems
turn yellow prior to dying. Picking the bulbils from the stem would result in getting rot-
ten ones even before they could be planted. Bulbils are commonly planted in September.
The porang produces an average of 50 bulbils per plant with, on average, a 3 cm diameter
(see Figure 4.2). The larger the bulbil diameter, the larger tuber diameter that will be har-
vested. However, farmers still pick out the bulbils with a minimum of 1 cm diameter for
replanting. The common range of the bulbil diameter is 1–5 cm. Prior to planting, bulbils
are kept in the airy, dry, and cool place on a rack.
FIGURE 4.2
Bulbils of A. muelleri.
The number of bulbils produced can be estimated from the number of the konjac planted
per ha, which is around 30,000 plants. The harvest ranges from 10,000 to 15,000 plants per
ha, and the bulbils are collected from the harvested mature plants. This means that the
production of bulbils is 500,000–750,000 per ha per year. Bulbils are sold on the market at
the price of IDR 40,000–50,000 (USD 3–3.5)/kg, and 1 kg of bulbils contains 80–100 pieces.
4.2.2.3.2 Replanting by Seeds

Seeds are collected from the konjac spadix. They are commonly harvested in August,
when part of the seeds turns red. One spadix could contain around 250 seeds. The spadi-
ces are kept leaning against the rack in the barn until all seeds turn red in about one week
indicating the physiological maturity.
The seeds are than removed from the spadix, then washed to separate the outer skin
layer. Bad seeds will float in the water and be separated from the good seeds. The good
seeds will be taken to dry in the ambient air on the bamboo mattress, and kept in well
ventilated, dry, and cool conditions on the rack.
When germinated, usually in September, the seeds will be planted in a polyethylene bag
until they grow to a plantlet of about 10 cm in length and have four leaflets when they are
taken to the field for planting.
4.2.3 Socio-Economic Aspects of Agroforestry

The farmers’ community living in the surrounding area of the KPH earns a good income
by selling the fresh tubers, sundried or mechanically dried konjac chips, and bulbils.
The price of fresh tubers in early 2018 was IDR 4–5 Million (USD 300–350) per tonne, of
dried chips IDR 30–35 Million (USD 2,000–2,350) per tonne, while for bulbils it was IDR
40,000–50,000 (USD 3–3.5) per kg, and 1 kg of bulbils contains 80–100 pieces with different
diameters. The conversion from fresh tubers to the dried konjac chips is 1/6. The average
yield of fresh tubers is 8 tonnes per ha ranging from 7 to 10 tonnes per ha. The average
konjac area is 1 ha per household. Individual farmer marks the border of their cultivated
field by growing turmeric and ginger, so that he can earn more income from them.
With the significant gain of income from the konjac industry, a lot of households can
live above the poverty line, and more importantly, they can support a better education for
their children. For example, Mr. Hartojo, the leader of the farmer community at Klangon
Village for more than 30 years could see all his three children enter and graduate from the
university.
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5
Postharvest Technology of Konjac
Rarisara Impaprasert, Zhao Jianrong, George Srzednicki, Yu Lei, and Tao Ruixuan
CONTENTS
5.1 Harvest and Delivery of Tubers to Processing Plants................................................... 161
5.1.1 Harvesting and Storage of Corms........................................................................ 161
5.1.2 Delivery to the Processing Plant........................................................................... 162
5.2 Primary Processing of Konjac Corms.............................................................................. 162
5.3 Browning Control............................................................................................................... 163
5.3.1 Browning Reaction................................................................................................. 163
5.3.2 Browning of Konjac Corms................................................................................... 164
5.3.3 Control of Enzymatic Browning Reaction.......................................................... 164
5.3.4 Browning Control of Konjac Corms..................................................................... 166
5.4 Drying and Storage of Dried Chips................................................................................. 167
References...................................................................................................................................... 169
5.1 Harvest and Delivery of Tubers to Processing Plants

5.1.1 Harvesting and Storage of Corms
In China and Japan, A. konjac is cultivated for food and is harvested 1 year after planting,
when the tubers are small, but sweet and juicy. For industrial purposes, the tubers are
harvested after three years. A single A. konjac tuber weighs about 200 g. after one year and
1–2 kg. after three years. Individual A. muelleri tubers may weigh up to 3 kg. depending on
the number of growing seasons. The A. muelleri propagated from small tubers is harvested
2.5 years after planting, whereas plants raised from bulbils are already harvestable after
1.5 years. A single A. variabilis tuber can weigh up to 1.5 kg.
The harvesting occurs at the end of the growing season, when the leaves begin to wither
and die. The konjac corms are carefully dug by hand because damage to them will facili-
tate infection by diseases and pests. The corms are usually well cleaned and are stored
in heaps, preferably in well ventilated sheds. They can lose as much as 25% of their initial
weight in the first month of storage, but can be successfully stored at 10°C for several
months. Alternatively, they may be left in the ground until required, with a little irriga-
tion if necessary. Corms dipped for one minute in a 4% fungicidal emulsion can be stored
at room temperature for about two months with minimal loss of weight or sprouting.
In Japan, the corms of the A. konjac that are to be replanted must be protected in storage
from low winter temperatures, since it has been found that if they are subjected to tem-
peratures of –5°C, germination is affected.
161
5.1.2 Delivery to the Processing Plant

Harvested tubers are brought to a yard near a road and cleaned first by removing the
earth, roots, and leaf parts. Then, they may be placed in heaps until they can be loaded
onto carts or trucks and transported to the processing plant. In order to obtain the best
quality of Konjac glucomannan (KGM), it is essential that they be transported to the pro-
cessing plant without delay. After the delivery at the processing plant, they will first be
peeled or grated, and the eyes (buds) and any rotten parts removed. Then they are washed
several times in water, graded according to the diameter of the corm, and sliced. For the
maximum production of KGM from the A. konjac or A. muelleri, the sliced tubers need to be
quickly dried first because within 24 hours the glucomannan breaks down enzymatically
in the presence of moisture.
5.2 Primary Processing of Konjac Corms

The primary processing includes all the operations (see Figure 5.1) that are performed
on the fresh corms. They constitute the first part of the dry processing as described in
Chapter 6. These operations may all be performed on a farm using traditional man-
ual methods and sun drying. However, the need to produce high-quality konjac flour
and the often large amount of freshly harvested corms make mechanized processing
unavoidable.
One of the operations that is generally mechanized, irrespective of the size of the enter-
prise, is washing, as it has to be performed thoroughly and rapidly. Figure 5.2 shows a
common type of equipment used for this purpose.
FIGURE 5.1
Flowchart of the primary postharvest operations.
Postharvest Technology of Konjac 163
FIGURE 5.2
Washing unit for konjac corms equipped with rotating brushes (George Srzednicki).
Cleaning and peeling can be done manually or mechanically in order to remove the top
buds, roots, and epidermis. However, the weak point is that the removed peel is about
1–2 cm. thick, which means about 3% of the KGM in the external cell layers is also dis-
carded. Should the sliced tubers be sun dried for more than 10 days, it will be more dif-
ficult to scrap the epidermis, and more layers of peripheral cells will have to be removed.
Therefore, the ‘non-scrapping washing method’ has been proposed for the wet pro-
cessing (described in Chapter 6). This method is based on the principle that the exter-
nal cell layers of the tuber don’t contain KGM, but are composed mostly of fibre tissue
(Li 2006). Once the corm tissue has been ground, some of the surface fibre is sieved into
waste liquid, and the rest is dehydrated and ground along with flour granules. During
drying, the surface fibre shrinks and, being much lighter than flour granules, it can be
easily separated from the granules in the air classification process. Hence, the quality
of the purified flour is not compromised, and all the KGM is recovered. The peels and
other residues left from cleaning and washing the fresh tubers can be used as fertiliser
for the nearby fields.
5.3 Browning Control

5.3.1 Browning Reaction
Browning is one of the most important reactions occurring during the food processing
and storage. The major groups of reactions leading to browning are the enzymatic phe-
nol oxidation and non-enzymatic browning reaction. Both of them can affect the quality
of food in either positive or negative ways, depending on the type of food (Manzocco
et al. 2001). Browning in fruits, vegetables, and seafood is often found due to the reaction
of polyphenol oxidase with phenolic compounds in the presence of oxygen and under
optimum conditions (Walker & Ferrar 1998). The phenolic compounds are oxidised to qui-
nones and changed to a brown substance. This reaction is called the enzymatic brown-
ing reaction. Meanwhile, the non-enzymatic-induced brown pigment formation, known
as the Maillard reaction, is caused by free amino groups, which are found in free amino
acids, peptides, or proteins and carbonyl groups, which are found in reducing sugars.
Food browning can be also due to the degradation of carbohydrates also called the cara-
melization reaction or the pyrolysis of sugar due to heat treatment above the melting point
of the sugar under alkaline or acidic conditions. Ascorbic acid, its oxidation products, and
oxidized lipids can also react via the Maillard reaction.
5.3.2 Browning of Konjac Corms

The important properties that are used to evaluate the quality of konjac flour are gluco-
mannan content, viscosity, and the colour of the konjac flour. According to the Chinese
grading powder standard, the white konjac powder is what customers want (Liu et al.
2002). Therefore, the white colour of the konjac powder is preferred by the konjac mar-
ket. However, the problem is that konjac always easily turns to brown when the flesh is
wounded and in contact with oxygen. In the konjac processing industry, one important
step is to dry the konjac in order to use it for processing throughout the year. A high
temperature during the drying process of the moist konjac cut slices can also trigger a
browning reaction. Based on the possibilities above, it seems that the browning of the
konjac is probably caused by the activity of the polyphenol oxidase (PPO) enzyme due to
the browning observed immediately after the flesh of the konjac is sliced and exposed to
oxygen. Because there are some minor concentrations of other carbohydrates (e.g., mono-
and disaccharide), proteins, and lipids in the fresh konjac, the drying takes place below
100°C in which the caramelization, Maillard reaction, or lipid oxidation would be very
difficult to occur. Moreover, there are the studies which found that browning of the konjac
was caused by the activity of the PPO (Zhang et al. 2007a, 2007b). For this reason, it can be
concluded that the enzymatic browning reaction is the main factor to the browning prob-
lem in dried konjac slices.
5.3.3 Control of Enzymatic Browning Reaction

PPOs (Emulsifiable concentrate 1.10.3.1) are a group of copper enzymes, widely distrib-
uted phylogenetically from bacteria to mammals, that catalyze the oxidation of pheno-
lics to quinones, which produce brown pigments in the wounded tissues (Mayer 1987).
The mechanism of action proposed for PPOs is based on their capacity to oxidise phenolic
compounds. When the tissue is damaged, the rupture of plastids, the cellular compart-
ment where PPOs are located, leads to the enzyme coming into contact with the phenolic
compounds released by the rupture of the vacuole, the main storage organelle of these
compounds. The active site of the PPOs consists of two copper atoms, and the enzyme
catalyzes two different reactions in the presence of molecular oxygen: the hydroxylation
of monophenols (monophenolase activity) and the oxidation of o-diphenols to o-quinones
(diphenolase activity). This reaction is followed by the non-enzymatic polymerization of
the quinones, giving rise to melanin, pigments of high molecular mass and dark colour
(Queiroz et al. 2008). The pathway of this reactions is shown in Figure 5.3.
According to the literature, polyphenol oxidase activity can be regulated by the action
of anti-oxidant agents, enzyme inhibitors, acidulants, chelating agents, or complexing
agents (Özoglu & Bayindirli 2002). The other methods and technologies, such as thermal
FIGURE 5.3
Polyphenol oxidase pathway. (Adapted from Corzo-Martínez, M. et al., Browning reactions, in Food Biochemistry
and Food Processing, 2nd edn., Simpson, B.K., Nollet, L.M.L., Toldrá, F., Benjakul, S., Paliyath, G. and Hui, Y.H.
(eds.), John Wiley & Sons, Iowa, IA, 2012.)
processing, exclusion of oxygen, and anti-browning agents, have been studied to inhibit poly-
phenol oxidase-related enzymatic browning (Loaiza-Velarde et al. 2003). However, a common
approach for preventing enzymatic browning is the use of anti-browning agents (Arslan &
Doğan 2005). Sulphur-containing agents are found to be successful inorganic salts to con-
trol browning in fruits and vegetables since 1986 (Son et al. 2001). Many compounds contain
sulphites, called sulphiting agents, including sulphur dioxide, sodium sulphite, sodium and
potassium bisulphites, and metabisulphites. Walker (1977) proposed that the primary role of
sulphiting agents is to react with the pigment precursors (o-quinone) to produce stable and
colourless diphenols before they can undergo further reaction to form pigments. Ferrer et al.
(1989) proposed that bisulphate inhibition was due to the reaction of sulphites with interme-
diate quinones, resulting in the formation of sulphoquinones, which irreversibly inhibited
polyphenol oxidase, causing complete inactivation. On the other hand, Madero and Finne
(1982) proposed that bisulphite exerted a competitive inhibitory effect on polyphenol oxi-
dase, by binding a sulfhydryl group at the active site of the enzyme. The destruction of the
disulphide bonds may cause the enzymes to denature or lose their shape because enzymes
must have a specific three-dimensional structure to catalyze their biochemical reactions
(Fu et al. 2007). They are currently applied for the inhibition of the browning in potatoes,
mushrooms, apples, and other fruits and vegetables. Sulphite concentrations necessary for
controlling enzymatic browning vary widely in accordance with the food material and the
time required for the inhibition of the browning reaction (Taylor et al. 1986). Although sul-
phite treatment levels in foods vary widely between applications, the residue levels should
not usually exceed several hundred parts per million (ppm) (Sapers 1993; Taylor et al. 1986).
Moreover, the Joint Expert Committee on Food Additives of the World Health Organization
and the Food and Agriculture Organization recommended an acceptable sulphite daily intake
of sulphur dioxide in foodstuffs as 0.7 mg/kg of body weight (Queiroz et al. 2008). In addi-
tion, these compounds have been restricted by the Food and Drug Administration due to
the possibility of their associated potential hazards to human health, especially in asthmatic
patients (Simon 1998). Thus, the Codex General Standard for Food Additives online database
limited the maximum level use of a sulphiting agent to 50 mg/kg for starches, 200 mg/kg for
flours, 500 mg/kg for dried vegetables (including mushrooms and fungi, roots and tubers,
pulses and legumes, and Aloe vera), seaweeds, and nuts and seeds, and 1000 mg/kg for dried
fruits (Codex Alimentarius 2019).
Hence, several studies have been devoted to the non-sulphite anti-browning agents
(e.g., ascorbic acid, citric acid, and sodium chloride) in many fruits and vegetables. Natural
anti-browning agents were also tested. Honey (Chen et al. 2000; Gacche et al. 2009);
banana leaf extract either alone or in combination with ascorbic acid and 4-hexylresorcinol
(Kaur and Kapoor 2000); papaya latex extract (De Rigal et al. 2001); onion juice (Hosoda
and Iwahashi 2002); solutions containing citric acid, calcium chloride, and garlic
extract (Ihl et al. 2003); onion oil (Hosoda et al. 2003); onion extracts (Kim et al. 2005);
onion by-products, such as residues and surpluses (Roldan et al. 2008); rice bran extract
(Boonsiripiphat and Theerakulkait 2009); and chitosan (Martin-Diana et al. 2009) are some
examples of natural inhibitors of PPOs. Inorganic halides are also well known inhibitors of
polyphenol oxidases (Vámos-Vigyázó 1981). Janovitz-Klapp et al. (1990) found that sodium
fluoride was the most potent inhibitor of apple polyphenol oxidase, followed by sodium
chloride, sodium bromide, and sodium iodide. Sodium chloride and calcium chloride at
concentrations ranging between 2% and 4% (w/v) are the most commonly used in the food
industry for the inhibition of browning (Steiner & Rieth 1989). Polyphenol oxidase activity
was observed to decrease with increasing concentrations of sodium chloride for the peach
(Luh & Phithakpol 1972), eggplant, and avocado (Knapp 1965).
5.3.4 Browning Control of Konjac Corms

In order to inhibit the browning reaction of konjac corms while they were dried for safe
storage, various anti-browning agents were studied. A number of studies are related to the
browning control in the konjac. Shimizu and Shimahara (1973) studied the extraction pro-
cess of konjac flour and used the sodium or potassium salt of sulphurous or hyposulphu-
rous acid in an amount of 100–200 ppm as an anti-browning agent. Chua et al. (2010) used
sulphur dioxide as a bleaching agent to prevent the darkening of the konjac chips. Zhao
et al. (2010) studied the effects of using four SO2 releasing agents including Na2SO3, K2SO3,
Na2S2O5, and NaHSO3 at a concentration of 0.1%–0.4%, and ascorbic acid at a concentration
of 6% on the discoloration of konjac flour in the extraction process. They found that all
four inorganic anti-browning agents provided good protection against the discoloration
of konjac flour at a concentration of 0.25%. Moreover, at this concentration, the sulphurous
acid radical remained below 40 ppm, which complies with the Chinese food standards.
Furthermore, using 6% of ascorbic acid resulted in a non-discoloured, sulphur-free konjac
flour. Chua et al. (2012) studied the methodologies for the extraction of the KGM from the
corms of A. konjac and used 1% (w/v) sodium bisulphite for one minute as an anti-brown-
ing agent. The experimental results were satisfactory. In addition, the konjac flour manu-
facturers commonly use sulphur dioxide in sulphur fumigation during the drying process
of konjac chips. According to the information obtained from the industrial sources, it can
be concluded that a sulphating agent is commonly used as an anti-browning agent to pre-
vent browning in konjac flour production. Impaprasert (2013) had conducted a new study
on the prevention of browning in dried konjac. Sodium chloride, potassium hydroxide,
sodium hypochlorite, sodium borate, and two sulphur-containing compounds including
sodium metabisulphite and potassium aluminium sulphate were used in order to inhibit
the formation of browning in the konjac slices before drying. The result indicated that
among the anti-browning agents, sodium metabisulphite at 500 ppm, with ten minutes of
soaking time, was the most effective anti-browning agent to retard the browning reaction,
while ascorbic acid exhibited the least inhibition activity for protection from browning of
the dried konjac slices. Ascorbic acid is a highly water-soluble and moderately reducing
compound, acidic in nature, and forms neutral salts with bases. Ascorbic acid, also known
as vitamin C, and its derivatives have been widely used as an anti-browning agent for the
processing of fruits and vegetables. In addition, ascorbic acid also acts as an oxygen scav-
enger for the removal of molecular oxygen in polyphenol oxidase reactions. Polyphenol
oxidase inhibition by ascorbic acid has been attributed to the reduction of enzymatically
formed o-quinones to their precursor diphenols (Walker 1977). Ascorbic acid is, however,
irreversibly oxidised to dehydroascorbic acid during the reduction process, thus allowing
browning to occur when the reducing efficiency of ascorbic acid was depleted. When com-
pared with the sulphur-free compounds group, sodium chloride at 10,000 ppm, soaking
for 20 minutes, showed a clearly better ability to inhibit the browning reaction in dried
konjac slices when compared with sodium metabisulphite. Thus, soaking konjac slices in
sodium chloride solution before drying can be an alternative way to replace the use of
sulphiting agents.
5.4 Drying and Storage of Dried Chips

The final stage of the primary processing is drying. The quality of the semi-processed
materials obtained in the first phase of the dry process (chips) cannot be controlled since
the sliced corms are generally dried either on a sunning floor or on a heated brick bed.
The modern processing plants use vibrating-fluidized bed dryers (Figure 5.4) or belt dry-
ers (Figure 5.5). However, drying is performed often with direct heating by flue gases.
Since a different quality of coal is used as a fuel for the mechanical dryers, the combustion
residues contaminate the dried product. Only the most advanced fluidized bed dryers
use heat exchangers. Nowadays, the traditionally used coal is increasingly replaced by
biomass, such as rice husks or other processing residues.
In order to prevent discoloration, SO2 is used, resulting in a high sulphur content of
the chips and subsequently of the konjac flour. Besides having a high sulphur content,
FIGURE 5.4
Vibrating fluidised bed dryers (Zhao Jianrong).
FIGURE 5.5
Belt dryer (George Srzednicki).
konjac flour produced by the dry process is often characterized by a low viscosity (Wu and
Zhang 1994). As a result, most of the product resulting from the traditional dry process-
ing can only be used as third grade flour. Such a product often cannot meet the Chinese
Professional Standard for Konjac Flour NY/T494-2002 (Ministry of Agriculture P. R. China
2002), especially with regard to product colour.
In order to obtain good physical and chemical properties of konjac flour, the processor
has to prevent it from browning, case hardening, and gelatinization (Huang 1994). During
processing, browning and case hardening would affect the colour of konjac chips, while
gelatinization would lead to the reduction of viscosity. At present, the traditionally dried
chips often have such defects: The sun-dried chips are often discoloured and have a low
viscosity. The mechanically dried chips may have acceptable colour, but often have exces-
sive sulphur content.
The konjac chips are generally packed in woven polyethylene bags and stored at or near
the drying plant. The processor may keep them in storage at an ambient temperature and
with scarce light for several months until they can be sold for further processing. It is
important that they be stored in a dry place to prevent moisture reuptake. If the plant has
the necessary equipment, the chips may be ground into powder which is called ‘common
konjac flour’, then sifted and separated from impurities. This product is called ‘common
konjac fine flour’. These products will also be packed in bags and kept in storage under
similar conditions as konjac chips.
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6
Processing of Konjac Flour
Rarisara Impaprasert, Zhao Jianrong, and George Srzednicki
CONTENTS
6.1 Principles of Konjac Processing........................................................................................ 173
6.2 Dry Processing.................................................................................................................... 175
6.2.1 Grinding and Sifting.............................................................................................. 175
6.3 Wet Processing.................................................................................................................... 177
6.4 Combined Wet and Dry Processing................................................................................. 178
6.4.1 Process Description of the Wet Processing Step................................................ 180
6.4.1.1 Determination of the Reaction Time..................................................... 180
6.4.1.2 Pre-Treatment in Anti-Swelling Solution............................................. 180
6.4.1.3 Pre-Treatment in Anti-Browning Solution........................................... 180
6.4.1.4 Crushing and Grinding.......................................................................... 181
6.4.1.5 Washing and Dehydration...................................................................... 181
6.4.1.6 Drying of Common Fine Flour.............................................................. 181
6.4.1.7 Recycling of Ethanol................................................................................ 182
6.4.2 Process Description of the Dry Processing Step................................................ 183
6.4.2.1 Grinding and Separation of Particles.................................................... 183
6.4.3 Packaging................................................................................................................. 183
6.5 Quality Control and Storage of Konjac Flour................................................................. 184
References...................................................................................................................................... 187
6.1 Principles of Konjac Processing

Konjac flour (gum) is an intermediate product that is a crude or refined product of gluco-
mannan. Konjac flour processing is the basis and key to the use of konjac, and its quality is
directly related to its application range, which includes food, medicine, chemical, textile,
petroleum, and other industries. According to early sources, the most primitive konjac
flour processing was invented by farmers in Ibaraki Prefecture, Japan. Nakajima Taniwa
(1745–1826) cut the konjac tubers into thin slices, dried them, and then ground them into
powder (coarse powder). Later on, a blower was installed on the grinder of the coarse
powder to blow away the fine particles from the powder. Since the middle of the twenti-
eth century, Japanese articles and patents concerning the processing of konjac flour have
appeared and new processing equipment for konjac was successively launched to perform
grinding and sifting operations. This equipment was originally developed for dry process-
ing. In order to improve the quality of the konjac flour a ‘wet method’ (extraction of gluco-
mannan from fresh konjac tubers through liquid media) was developed.
The different grades of Konjac Flour (KF) are listed in Table 6.1. Table 6.1 shows that
KF is divided into the four categories according to NY/Y 494-2002 ‘China’s Agricultural
173
TABLE 6.1
Definitions of Different Types of Konjac Flour
Name of Processing Product
Product Raw Materials Processing Method Characteristics Characteristics
Common konjac Konjac chips (bar) Physical dry processing Starch removal Particle size: 90%
fine flour Konjac fresh tuber Food grade ethanol wet processing 0.125–0.335 mm
Common konjac Konjac chips (bar) Physical dry processing Starch removal Particle size: 90%
particulate flour Konjac fresh tuber Food grade ethanol wet processing ≤0.125 mm
Purified konjac Konjac fresh tuber Food grade ethanol wet processing Purify & extract Particle size: 90%
fine flour Konjac flour Food grade ethanol purification KGM 0.125–0.335 mm
KGM% ≥ 85%
Purified konjac Konjac fresh tuber Food grade ethanol wet processing Purify & extract Particle size: 90%
particulate flour Konjac flour Food grade ethanol purification KGM ≤0.125 mm
KGM% ≥ 85%
Standard for Konjac’ (Ministry of Agriculture P.R.C. 2002). Common konjac particulate
flour is a semi-finished product that through further processing may become a refined
product (Chong 1990). The processing of konjac tubers into flour is the key step in the uti-
lization of konjac. The quality of the final product determines its range of applications, i.e.,
pharmaceuticals, food, chemicals, textile, or oil industries (Matsuzaki 1992).
The focus of konjac processing is on the extraction of Konjac Glucomannan (KGM) from
corms (Chen and Sun 1989). The processing methods can be divided into three catego-
ries depending on whether and when liquid media are used in the process, as shown in
Figure 6.1 (Iwatani 1978; Wootton et al. 1993; Sun and Yang 1994).
FIGURE 6.1
Processing methods of KF. (Adapted from Iwatani, S.K., Powdered konjac product, J88065297, J55092667, 1978;
Wootton, A.N. et al., J. Sci. Food Agric., 61, 429–433, 1993; Sun, Y. and Yang, Y., The processing technology of
konjac flour, ZL94116535.3, 1994.)
Processing of Konjac Flour 175
FIGURE 6.2
Flowchart showing dry processing of konjac corms.
6.2 Dry Processing

The process starts by slicing and drying konjac tubers. The dried chips are ground into
powder which is called ‘common konjac flour’. The latter is then sifted and separated from
any impurities. This product is called ‘common konjac fine flour’. The dry processing
method involves two parts: the first is to produce ‘common konjac flour’ and the second is
to further grade it to ‘common konjac fine flour’. Since the whole process only involves dry-
ing and mechanical separation it is called ‘dry processing’. Figure 6.2 shows the schematic
of the dry processing of fresh corms.
6.2.1 Grinding and Sifting

The equipment required for grinding of the dried chips and sifting of the coarse powder
are shown in Figures 6.3 and 6.4.
FIGURE 6.3
Grinder for konjac chips (George Srzednicki).
FIGURE 6.4
Sifter for separation of different fractions of konjac flour (George Srzednicki).
6.3 Wet Processing

Wet processing involves using a protective solution for dipping the freshly crushed tubers.
As a result, the konjac flour does not swell or turn brown. Wet processing involves crush-
ing, grinding, centrifugation, and drying. The traditional wet processing technique may
be performed in two ways as shown as Figures 6.5 and 6.6. The first one consists of pro-
ducing KF directly from fresh tubers, whereas the other one uses ‘common konjac flour’
obtained from dry processing in order to purify it (Huang 1997, 2005). The cost of the
former is high, while the quality of the KF obtained by the latter process is not uniform.
The protective solution used in the wet process may use organic or inorganic solutions.
The organic solution uses generally food grade ethanol as a solvent, whereas the most
common constituent of the inorganic solution is 3‰ borax (Na2B4O7 · 10 H2O) (Zhang 1994).
The KF produced in the former way is of good quality and can be used for various applica-
tions, such as pharmaceuticals and food, but the cost is high. The latter costs less, but the
KF isn’t edible and cannot be used as food.
FIGURE 6.5
Flowchart of wet-processing using fresh corms as raw material.
FIGURE 6.6
Flowchart of wet-processing using common KF as raw material.
6.4 Combined Wet and Dry Processing

In order to satisfy the growing demand of the market, the concept of producing purified
KF characterized by high output and high viscosity with low sulphur content and low
production cost has been devised. By combining the elements of wet and dry processing
in a two-stage technique (see Figure 6.7), the total output can be increased, whereas the
consumption of the protective solution and energy can be reduced.
The process is based on the following principle: KGM and starch are the main two ingre-
dients of konjac fresh corms and need to be separated efficiently without affecting the
quality of the KGM. During their short stay in the protective solution, the idioblasts remain
hard and tough and are not easy to crash by the mechanical force of shearing or impact.
In contrast, the ordinary cell can easily be crashed into smaller particles because of its
low hardness and high brittleness. Meanwhile, during the contact between the konjac and
FIGURE 6.7
Flowchart of the combined wet and dry process.
liquid medium, some soluble impurities (e.g., some oligosaccharides and alkaloids) of the
KGM idioblast are dissolved gradually. Then, during the solid-liquid separation, the starch
dust will be filtered through filter cloth with a certain aperture size and soluble impurities
will be removed in the waste solution. At this moment, the first processing stage, the wet-
processing, is finished. In the second stage, the dry processing, the surface impurities of
the KGM idioblast will be removed constantly through the impact and friction between the
KGM granules and machine, and then separated by sifting, as shown in Figure 6.7.
Figure 6.7 shows that the first step consists of immersing the crushed tubers (but without
epidermis or subepidermal tissue) into a series of organic and inorganic solutions at a specific
concentration at which the macromolecules can expand without dissolving and becoming
mushy. The macromolecules can be broken up gradually and efficiently by a disintegrator and
their components separated. This is the wet processing stage, during which, the KGM gran-
ules will be purified by removing alkaloids, tannins, starch, sulphur, and other impurities.
The second step, the dry processing stage, consists of grinding and sifting the KF obtained
during the first step following the specifications of the application in which it will be used.
A further separation of starch from the surface of the KGM granules occurs at this stage.
In brief, the first step is to purify the rough KF by the wet process, whereas the second
step is to separate the KF obtained from the first step by the dry process. The KF produced
by this two-stage technique is of good quality like that produced directly by wet pro-
cessing, but the production cost is much lower. At a large industrial scale, this technique
requires high-speed and high-capacity processing equipment for crushing raw tubers,
grinding and sifting granular material, and drying and recycling ethanol (Huang 2000;
Ishii and Kasuga 1991).
6.4.1 Process Description of the Wet Processing Step

The first stage is also called wet milling, as all operations leading to the production of
purified rough flour are dealing with high moisture material.
6.4.1.1 Determination of the Reaction Time

Since glucomannan is a hydrocolloid, it is very important to avoid the swelling of the KGM
in the water-ethanol solution during the wet process in order to minimize the production
cost. It had been established (Liu 2004; Huang 1997) that there is a time limit within which
the KGM in A. muelleri, A. bulbifer, and A. konjac will be insoluble in a 55% water-ethanol
solution. This time was experimentally determined in order to define the safe time for
corm processing during the wet process stage.
6.4.1.2 Pre-Treatment in Anti-Swelling Solution

The protective solution is composed of water and anti-swelling and anti-browning agents
in well determined proportions. It protects the KGM from expanding in water, becoming
brown and black, and assures normal operation of equipment.
Because the KGM is highly hydrophilic, it easily dissolves in water and a sol is formed.
Therefore, ethanol is used in different concentrations in the wet process in order to control
the solubility of the KGM in the water-ethanol solution. The composition of the KGM var-
ies in different species and so does its solubility in the water-ethanol solution.
In practice, the ethanol concentration will be decreasing with an increasing weight ratio
(ethanol:fresh tubers) which is between 1 and 3 (Huang 2005; Ai and Ai 2000).
If the ratio is 1 to 1, the ethanol concentration can be above 70%.

If the ratio is 3 to 1, the concentration can be above 35%.
If the ratio is 2 to 1, the concentration can be 50% to 60%.
6.4.1.3 Pre-Treatment in Anti-Browning Solution

Because the konjac tissue can become dense paste when in contact with water and because
corms contain pigments, the paste may become discoloured. The impurities that cause dis-
coloration are difficult to remove from the konjac paste, resulting in a reduction of quality
of the purified KF. To avoid this problem, anti-browning agents are used.
Sulphites are generally applied for the inhibition of the browning in fruits and vegeta-
bles. Sulphite concentrations necessary for controlling enzymatic browning vary widely
in accordance with the food material and the time required for inhibition of the browning
reaction (Taylor et al. 1986). However, these compounds have been restricted by the Food
and Drug Administration (FDA) due to the possibility of associated potential hazards to
human health.
6.4.1.4 Crushing and Grinding

This process takes place in the protective solution. It involves a grinding mill followed by
a colloid mill.
A grinding mill is operated at a high rotational speed, producing a strong shearing and
grinding of the material. The material in the protective solution is forced into a cavity
formed by a spinning rotor and fixed stator. A centrifugal force propels the material to the
outside of the rotor, causing intense hydraulic shear that breaks agglomerates and homog-
enizes the solids and liquids.
The second machine is a colloid mill. It is a device that is reducing the particle size of a
solid in suspension in a liquid by applying high levels of hydraulic shear to the process
liquid. It is generally used to increase the stability of suspensions and emulsions. High
rotational speeds, combined with an extremely small shear gap, produce intense friction
on the material being processed. The rotor and stator are cone shaped and have increas-
ingly fine serrations or grooves. The friction and shear that result are commonly referred
to as wet milling.
6.4.1.5 Washing and Dehydration

The supernatant liquid above the ground mass can be used as a washing liquid. Washing
and dehydration can be operated simultaneously. Dehydrating is carried out in a centri-
fuge fitted with a screen with 149 μm (100 mesh) openings. During the operation of the
centrifuge, the washing water is poured slowly and continuously to the central centrifuge
spindle. This operation is carried out at a high rotating speed until the effluent liquid is
almost clear from centrifuge. Thus, the impact force of the washing water is increased, and
the washing effect is better. After leaving the centrifuge, the supernatant liquid with an
ethanol content of about 30% flows into the waste sedimentation tank. After sedimentation
of the suspended solid particles, the washing water can be re-used. Compared with the
traditional wet process, the use of a centrifuge is not only avoiding repetitive washing in
the protective agent, but also reducing the time of the operation and reducing the cost of
materials.
The liquid with suspended sediment will be drained into a waste pool. The water con-
tent of the pulp after dehydration is 40% to 60%. Normally, due to the thorough action of
the grinding and of the colloid mill, the KF is free of large agglomerated granules, as they
are in a dispersed state. The process of crushing and grinding of the A. bulbifer is oper-
ated in water (acting as an anti-swelling agent), as long as it does not exceed 45 minutes.
Beyond this time, the KGM particles may start swelling. Should the process last longer than
45 minutes, the washing liquid should include 30% ethanol in order to prevent swelling.
6.4.1.6 Drying of Common Fine Flour

Drying is one of the key procedures in the wet processing, as it determines the final qual-
ity of the KF. Given the hygroscopic nature of the KGM, it has to be dehydrated without
delay. However, a high drying temperature can significantly reduce the viscosity of the KF.
The two most commonly used drying methods in wet processing, hot-air (pneu-
matic dryer) and vacuum drying, have their respective advantages and disadvantages.
The advantages of hot air drying are high speed (a few seconds) and high efficiency.
The inlet air temperature in the hot-air method is 120°C–150°C. Hence, a short drying
time is sufficient, and the quality of purified flour is not affected. The disadvantages of the
hot-air drying method are that the ethanol gas is not easy to collect and leaves a linger-
ing ethanol smell. As for vacuum drying method, the advantages are that ethanol vapour
is easy to recycle. In contrast, both process speed and efficiency are low. The long drying
time makes the KF darker and its viscosity lower. Therefore, vacuum drying cannot be
applied as a single drying stage.
In the first stage, the particles of the KF can freely flow in the drying air, while the
water-ethanol solution is rapidly vaporizing. In the hot air, the particles of the fresh
KF are dispersed into granular powder and transported in the stream of hot air. Wet
particulate materials that are dispersed and suspended in hot air flowing at a high
speed are subjected to a high rate of heat and mass transfer between the wet materials
and hot air. In the process, water and ethanol evaporate, the particulate materials enter
a cyclone separator and are collected at the bottom of the machine, while the exhaust
gases are vented. Because the airspeed in the pneumatic dryer is 20 ~ 30 m/s, the con-
tact time of the gas-solid is very short (about two seconds), so the powder is not likely
to be overdried.
In the second stage, the particles of the fresh KF are constantly moving, following the
rotation of the drying chamber. The heat transfer occurs through the internal surface of
the drying chamber, which is in direct contact with the particulate materials. Water and
ethanol will evaporate and be exhausted by a vacuum pump. In this process, due to the
action of the vacuum and uniform heating, water and ethanol will be removed completely.
A condenser is used for ethanol recycling.
Thus, in order to get the optimum results (complete recycling of ethanol, disappearance
of ethanol smell, and high efficiency), a combination of hot-air and vacuum drying are
recommended. The hot air removes the moisture quickly, resulting in the KF with a final
moisture content of about 10% wet basis; the vacuum dryer slowly removes the ethanol
vapours and smell from the product.
6.4.1.7 Recycling of Ethanol

In the wet process, the recycling of ethanol consists of the distillation of waste liquid so
that the ethanol can be re-used in the wet process. The conditions in the recycler are as
follows: the temperature at the bottom of the tower is 95°C–105°C, the distillation tempera-
ture is 78°C–88°C, and the vapour pressure is 0.08–0.12 MPa. Under these conditions, the
recycling rate of ethanol is at least 97%. The waste liquid left from the distillation should
be drained into a waste pool to be used for other applications.
In the process, ethanol recycle includes the following two parts (see Figure 6.8):
1. In the course of grinding and dehydration: there is a large volume of waste

liquid containing water, ethanol, ash, alkaloid, low grade fine KF, and impuri-
ties. Using distillation, some components of the waste liquid will be vaporized;
resulting in a separation according to the different boiling points (BP) and the
volatility of different components (BP of ethanol: 78°C; BP of water: 100°C).
This separation is performed by mass transfer between the liquid phase and
vapour phase.
FIGURE 6.8
Flowchart of ethanol recycling.
2. In the course of vacuum drying: the principle of ethanol recycle is the same as
mentioned above. A condenser is connected to the exit of the exhaust gas for water
and ethanol vapour liquefaction. This liquid will be collected and recycled like
that from the waste water.
6.4.2 Process Description of the Dry Processing Step

The second stage, dry milling, deals with the mechanical separation of the KGM resulting
in the production of purified fine flour.
6.4.2.1 Grinding and Separation of Particles

The principle of dry processing is as follows: konjac flakes are mainly composed of KGM
and starch granules that will be separated mechanically. The former accounts for about
55% and the latter for the remainder of the weight. The two types of granules differ in
size and hardness and thus can be separated mechanically. The high-speed grinders first
break up the softer starch granules and fibre cells (coming from peripheral cell layers of
the tubers). These granules will become disintegrated into tiny, ash-like, particles. During
this process, the KGM granules will generally not be broken up, as they are harder. Due to
the difference in size and weight, the KGM can be separated from the starch particles by
sifting or elutriation. Processing facilities for producing purified fine konjac flour include
grinders, cyclones, and sieves.
6.4.3 Packaging
A double-layer packaging is normally applied for the KF. The inner layer is a polyethyl-
ene film, and the outer layer consists of a woven polyethylene bag, paper, or laminate.
The product is kept in a dry and damp-proof enclosure without being exposed to sun-
shine. The optimum temperature should be below 25°C, and the relative humidity below
65%. If it is stored separately from other goods, the shelf life should be up to two years.
6.5 Quality Control and Storage of Konjac Flour

Nowadays, there are many types of powdered foods. The powder food industries have
grown continuously for many years since the convenience of eating and storage is what
consumers want. Powdered food, therefore, responds to all aspects, because of the conve-
nience in portability, preparation, and long storage, even without a refrigerator. Powdered
food has free water content (water activity) below 0.6 or humidity less than 15%, so it does
not support the growth of spoilage microbes including moulds, yeasts, and bacteria (Jay
1998). Powdered food can also be designed with complete nutritional value. Examples of
currently available powdered foods are milk, coffee, tea, cocoa, fruit juice, herbal juice,
soup, sugar congee, seasonings, vegetables, spices, and flour. However, the most common
problem with powdered food is caking, which is due to the fact that the components in
the food absorb moisture from the surrounding environment, causing the food to change
both physical and chemical properties. Changes in humidity during storage are impor-
tant factors for the quality and stability of powdered food. Storage conditions that directly
affect the quality of food include storage temperature and surrounding relative humidity.
In addition, each food has a different ability to absorb and release moisture. Therefore,
studying the proper storage conditions for each food type is important. This is because
the data can be used to control the quality of the product during storage and to predict the
shelf life of the product. The benefit of studying the optimum conditions is that this can
make it possible to select the suitable storage conditions and packaging for each food.
The hygroscopic character of food is connected with its ability to absorb water in a humid
environment or to release water in a dry environment (Domian & Lenart 2000). The com-
ponents in food that are hygroscopic are salts, polysaccharides, and proteins, since they
can form bonds with water. Water can create ionic bonds with salt ions, hydrogen bonds
with hydroxyl groups of sugar in polysaccharides, and carbonyl groups of amino acids
in proteins. Water that forms strong bonds with food components is called bound water,
which is a very small proportion of the total water content in a food. The water that is
weakly bonded with the components in food is called free water, which has a high propor-
tion compared to the total amount of water in food (Damodaran et al. 2008). Germination
improves the water adsorption ability of starch and protein-based flour, e.g., chickpea,
lentil, and yellow pea flour because of the structural changes of the macronutrients during
germination. Starch in the germinated pulse flour tended to transform from their semi-
crystalline form into an amorphous form, while proteins in the endosperm were loosened
so that a larger surface area was available to interact with the water molecules. Thus, the
increased water adsorption isotherm implied that more ionic interaction and hydrogen
bonding occurred between constituents in pulse flour and water molecules (Xu et al. 2019).
A reliable method to study the moisture changes in food during storage for food sta-
bility is the study of moisture sorption isotherm. The moisture sorption isotherm of a
material represents the equilibrium relationship between the water activity (aw) and its
moisture content at constant temperature and pressure. This relationship is complex
and unique for each product (Damodaran et al. 2008). The sorption isotherms are neces-
sary to predict the properties of the material in different environments pertinent to their
applications. Brunauer et al. (1940) classified sorption isotherms into five types includ-
ing: type I isotherms (Langmuir isotherms), for the monolayer adsorption that occurs
in porous solids; type II isotherms, for common soluble products, powder, and starchy
foods with a sigmoidal shaped curve (Basu et al. 2006; Ertugay & Certel 2000); type III iso-
therms (Flory-Higgins isotherms), for solvents and plasticizers above the glass transition
temperature; type IV isotherms, for hydrophilic solids with swelling properties in which
adsorption continues until the maximum hydration of sites; and type V isotherms, for
mesoporous materials, such as charcoal (Basu et al. 2006; Ozturk & Takhar 2018). There are
many studies about moisture sorption isotherm as follows:
Palich and Ruszkowska (2007) studied the hygroscopicity of food concentrates with
the example of pea soup. The sorption isotherms were determined with a static
method. Samples were stored in hygrostats at a temperature of 20°C ± 1°C, with
saturated saline solutions with a water activity from 0.07 to 0.98 for 30 and 90 days
for croutons and powder samples, respectively. The results from this research
indicated that the sorption isotherms of croutons are corresponding to type II,
while the sorption isotherms for pea soup concentrate powder were the indirect
type between II and III according to the Brunauer’s classification.
Ozturk and Takhar (2018) reviewed the water transport in starchy foods and the
changes occurring in the functional properties, such as moisture sorption, swell-
ing, gelatinization, and glass transition characteristics of starchy foods. Starchy
foods presented type II (sigmoidal shaped) isotherms, which indicated that they
underwent multilayer adsorption.
Kruangam and Intipunya (2013) studied the sorption isotherm and quality changes
during the storage of a powdered longan cube. The results indicated that the sorp-
tion isotherm of the sample was adsorption isotherm. The moisture content of
the sample increased as the relative humidity increased. It was also found that
the longan cube was sensitive to the adsorption of water from the environment.
There were significant quality changes during storage including colour, solubil-
ity, water activity, and moisture content. The samples became gel-like and had a
darker colour when stored in a high relative humidity condition. The researchers
suggested that the longan cube should be packed in a low moisture permeable
packaging and stored at a relative humidity of 32%–43%.
Moisture sorption isotherms were also successfully studied using a static gravimetric
method for nixtamalized amaranth flour (Valdez-Niebla et al. 1993); maize flour (Oyelade
et al. 2008); chestnut and wheat flours (Moreira et al. 2010); ahipa and cassava flours and
starches (Doporto et al. 2012); tef (Eragrostis tef) grain flours (Abebe & Ronda 2015); spray-
dried pure orange juice powder (Sormoli & Langrish 2014); spray-dried tamarind pulp
powder (Muzaffar & Kumar 2016); by-products from the rice industry (Torres & Seijo 2016);
cassava bagasse (Polachini et al. 2016); extruded snacks (Wani & Kumar 2016); raw and
extruded whole meal sorghum flours (Galdeano et al. 2018); and chickpea, lentil, and yel-
low pea flours (Xu et al. 2019). According to the Brunauer, Emmett, and Teller classifica-
tion, the isotherms for all systems are type II isotherms (a sigmoidal shape isotherm),
which is generally observed in complex foods containing proteins and polysaccharides.
This type of isotherm can be classified into three regions: the first one for the monolayer
moisture, which is strongly bounded into the product matrix; the second is almost lin-
ear, corresponding to the multilayered water; and the third region is related to the free
water available for chemical reactions. These isotherms are influenced by temperature.
The Guggenheim, Anderson, and de Boer model nicely fit and gave the best result for all
experimental isotherms.
The structures of food materials are very complex, so the prediction of equilibrium
moisture content is difficult and can cause higher variations (Zapata et al. 2014). Thus,
the mathematical models are used to correlate with experimental data to obtain semi-
empirical sorption isotherms. There are several models for describing the sorption behav-
iour. The Brunauer, Emmett, and Teller model is good at explaining multilayer sorption,
while the Guggenheim, Anderson, and de Boer model is known for its versatility (Zapata
et al. 2014). The Smith model is known for its ability to describe the sorption isotherms of
biopolymers like cellulose and starch (Smith 1947). The Halsey model provides a good rep-
resentation of the sorption behaviour of foods containing starch like corn flour (Andrade
et al. 2011). The Henderson model has been used for cereal grains (Zapata et al. 2014).
Konjac flour is a type of powder product, which has a polysaccharide base with hygro-
scopic properties to absorb water from the surrounding environment. Konjac powder is
very useful for both food and non-food industries, causing the world market to demand
large quantities of the konjac powder. Therefore, konjac powder manufacturers need to
increase the production capacity and stock a large quantity of konjac powder in order to
respond to the growing market demand. However, storing large quantities of konjac pow-
der in inappropriate conditions may affect its physical and chemical properties. Therefore,
it is necessary to study the optimum conditions for the storage of konjac flour. However,
no scientific evidence has been found regarding the suitable conditions for the storage of
konjac flour. Thus, the study of related research is important as a guideline for studying
the suitable conditions for konjac flour storage.
From the analysis of moisture absorption of konjac powder using the static gravimetric
method, it was found that the moisture content affects the moisture absorption of konjac
powder. The moisture content of konjac powder tends to increase steadily as the relative
humidity of the environment increases. The moisture absorption curve of konjac powder
is type II and called sigmoid or S-shape, which is a normal food (see Figure 6.9).
From the results of the experiment, it was found that the temperature, relative humid-
ity, and storage time affected the amount of free water of the konjac powder. A higher
temperature and relative humidity along with longer storage time resulted in the increase
in the amount of free water in the konjac powder due to the ability of konjac powder to
FIGURE 6.9
The moisture absorption curve of konjac powder.
absorb moisture from the surrounding air. However, when analysing the amount of free
water in konjac powder under the same relative humidity at different temperatures, it was
found that the konjac powder stored at high temperatures contained a lesser amount of
free water than konjac powder stored at low temperatures. This may suggest that a higher
temperature caused free water in the konjac powder to evaporate.
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7
Physico-Chemical Properties of Konjac
Glucomannan
Patricia Le Bail, Céline Lafarge, and Nathalie Cayot
CONTENTS
7.1 Structure of Konjac Glucomannan................................................................................... 189
7.1.1 Legislation in Europe............................................................................................. 190
7.1.2 Structure................................................................................................................... 190
7.1.3 Gelation Mechanism of Konjac............................................................................. 191
7.2 Physical Properties of Konjac Glucomannan.................................................................. 193
7.2.1 Water Solubility and Water Absorption.............................................................. 193
7.2.2 Thickening Agent................................................................................................... 195
7.2.3 Thermally Stable Gels............................................................................................ 197
7.3 Gelling Properties with Other Polysaccharides............................................................. 199
7.3.1 Starch-Konjac Mixed Systems............................................................................... 199
7.3.2 Encapsulation and Complexation......................................................................... 202
7.4 Structural Properties in Presence of Water and Cryoprotective Properties.............. 203
References...................................................................................................................................... 205
7.1 Structure of Konjac Glucomannan

The konjac flour is obtained from the tuber of the Amorphophallus konjac, a plant of the
Araceae family. This plant is found in the wild and cultivated as a vegetable in Thailand,
China, Vietnam, Korea, and Japan since the nineteenth century AD (Chua et al. 2012; Li
et al. 2005). The major compound of konjac flours is glucomannan, the reserve polysaccha-
ride of konjac. The glucomannans are also present under the same structural type in vari-
ous plant species: orchid bulbs, lily, and Aloe vera seeds (Dorthe 2005). The konjac flour is
obtained by crunching the tubers more or less finely. These are harvested after 2 or 3 years
of plant development. The tubers are cut, dried, crushed, and sieved. The resulting flour
contains between 51% and 72% dry weight of the konjac glucomannan (Fang and Wu 2004).
In order to obtain pure flour containing more than 95% of the konjac glucomannan in dry
mass, a purification step by hydroalcoholic washes is essential. Depending on the species,
the dried crude konjac flour contains about 10%–30% starch, 2%–5% fibre, 5%–14% protein,
3%–5% reducing sugars, and 3.4%–5.3% ash, it is low in vitamins and fat. The konjac flour
production accounts for 25,000 tonnes per year, with China and Japan as leading produc-
ers and consumers of the konjac flour. China has become the largest producer of the konjac
ahead of Japan, exporting more than half of its production. Japan retains its konjac produc-
tion for domestic consumption and has specialised in the production of pharmaceutical
grade flour for export, which is more limited in volume.
189
7.1.1 Legislation in Europe

The konjac glucomannan, which is exclusively derived from the species Amorphophallus
konjac, is authorised in Europe as a thickening additive under the number E 425 (ii) under
Directive 95/2/EC of the European Parliament and of the Council of 20 February 1995 on
food additives other than colorants and sweeteners. As an additive, the maximum specific
quantity is fixed at 10 g (single or mixture)/kg of foodstuff (European Union Regulation
No 1129/2011, Official Journal of the European Union L295/1).
The European Union Regulation (No. 231/2012, Official Journal of the European Union
L83/1) sets the specifications for the konjac glucomannan (Table 7.1).
7.1.2 Structure
The primary component in konjac flour from the Amorphophallus konjac species is kon-
jac glucomannan (KGM), a high molecular polysaccharide, see Figure 7.1. It is a slightly
branched polysaccharide having a molecular weight of 200,000–2,000,000 Daltons (actual
molecular weight of KGM depends on the processing method and storage time of the raw
material) (Yuan 2017).
The polysaccharide glucomannan consists of a linear arrangement of β-D-glucose (G) and
β-D-mannose (M) residues linked by 1–4 linkages. They occur in an approximate ratio of
5:8. The basic polymeric repeating unit has the following pattern: GGMMGMMMMMGGM.
Short side chains of 11–16 monosaccharides occur at intervals of approximately 50–60 units
of the main chain, attached by 1–3 linkages. Acetate groups at the C-6 position occur at every
9–19 units along the linear chain. The hydrolysis of the acetate groups favours the formation
of inter-chain hydrogen bonds, a feature which is partly responsible for the formation of gels
and its subsequent high water-absorption capacity and voluminous expansion (Dorthe 2005).
Dorthe studied, in 2005, 28 representative samples of the main species of the konjac
(Amorphophallus sp) in Asia. The flours were compared according to the konjac species and
the purification treatment. These studies have shown that regardless of the species, the
molecular structure and molar mass of the KGM raw flours are similar. The KGM struc-
ture varies only according to the flour purification process. The composition of raw flours
seems to be independent of the konjac species.
TABLE 7.1
European Union Specifications for Konjac Glucomannan
Criteria Specifications
Aspect Fine white to slightly brownish powder, fluid & odourless

Loss on drying <8% (105°C during 3 h)
Starch content <1%
Protein content <1% (N × 5,7)
Viscosity solution at 1% >20 Pa.s
Soluble matter in ether <0.5%
Soluble matter in alcohol at 50% <2%
Sulphite (expressed in SO2) <4 mg/kg
Chloride <0.02%
Lead <1 mg/kg
Total ash <2% (800°C, 3 à 4 h)
Physico-Chemical Properties of Konjac Glucomannan 191
FIGURE 7.1
Chemical structure of konjac glucomannan according. (From Zhang, C. et al., Carbohydr. Polym., 104, 175–181,
2014.)
A structural model of the konjac glucomannan (mannose:glucose ratio 1.8:1) has been
proposed based on X-ray data and stereo-chemical constraints. The unit cell dimensions
of the orthorhombic I222 space group are a = 9.01, b = 16.73, and c (fibre axis) = 10.40Å.
The structure crystallises in the mannan II polymorphic form in which the backbone
conformation of two-fold helices and the chains are oriented in an anti-parallel fashion
(Ogawa 1991).
As a gelling agent, the konjac glucomannan is quite unique in its ability to form ther-
moreversible and thermoirreversible gels under different and well controlled conditions.
The solutions of the native konjac mannan do not form a gel because the acetyl content pre-
vents the polysaccharide chains from approaching and interacting strongly. However, gels
are obtained when the solution is heated at a pH of 9–10. These gels will remain stable to
heat and under repeated heating up to 200°C. When exposed to a slight alkaline environ-
ment, the konjac solution forms thermoirreversible gels after cooling from the hot solution.
The non-reversibility of the gel occurs from the removal of the acetate groups, and partial
structural crystallization occurs due to the formation of inter-chain hydrogen bonds.
7.1.3 Gelation Mechanism of Konjac

The konjac glucomannan gels are obtained by heating in the presence of basic compounds
(pH between 11, 3, and 12.6) or large amounts of salts (K3PO4, Na3PO4) (Nishinari et al.
1992). The most commonly used basic compounds are: NaOH, KOH, Ca(OH)2, Na2CO3,
and K2CO3 (Zhang et al. 2014). The gels obtained are stable, firm, and thermoirreversible.
The exact gelling mechanism requires additional investigations, but at least two impor-
tant factors are known. The first factor is the hydrogen bonds. In the presence of an alkaline
agent, the konjac glucomannan molecule irreversibly loses the acetyl groups of its molecule.
This deacetylation facilitates the establishment of hydrogen bonds between konjac glu-
comannan chains, resulting in the formation of a structured gel network (Alonso-Sande
Teijeiro-Osorio et al. 2009). The second factor is the establishment of hydrophobic interac-
tions between konjac glucomannan molecules (Figure 7.2) (Wang et al. 2015). The lowest
critical concentration of the konjac glucomannan required to form an alkaline gel is esti-
mated at 0.5% (Nishinari et al. 1992).
The gelation occurs after a period of induction. The induction period is defined as the sum
of the deacetylation step and the aggregation step. The speed of the induction period would
be related to the concentration of the hydroxyl ions. When the speed is fast, both steps seem
indistinguishable (Yoshimura and Nishinari 1999; Williams et al. 2000; Dorthe 2005).
A number of parameters affect the gelation mechanism and, consequently, the proper-
ties of the final structure of the gel: the degree of acetylation, the temperature, the con-
centration and the molecular weight of the konjac glucomannan, and the concentration of
basic elements. The specific influences of each parameter are described in Table 7.2.
Nishinari et al. (1992) showed that the gels formed by the addition of salts had a lower
texture compared to the alkaline gels. The amount of salts required for the gel formation
is much higher than the amount of alkaline compounds. The authors also report that the
alkaline agent is not required to form a gel if the concentration of the glucomannan konjac
is greater than 8%.
FIGURE 7.2
Mechanism of gelation of konjac glucomannan. (Alonso-Sande, M. et al., Eur. J. Pharm. Biopharm., 72, 453–462,
2009.)
TABLE 7.2
Influence of Different Parameters on the Gelation Mechanism of Konjac Glucomannan
Evolution of the Parameters Helping the Formation of a Konjac
Glucomannan Gel Mechanism
↓ Acetylation degree ↑ Hydrogen bond formation

↑ Molar mass ↑ Linkage zones number
↑ Length of connected chains
↑ Konjac glucomannan concentration ↑ Molecule number
↑ Proximity between molecules
↑ Temperature ↑ Hydrogen bond formation
↑ Alkaline agent concentration ↑ Deacetylation
↑ Hydrogen bond formation
Source: Alonso-Sande, M. et al., Eur. J. Pharm Biopharm., 72, 453–462, 2009.
7.2 Physical Properties of Konjac Glucomannan

The KGM can form a highly viscous solution upon mixing and heating with water and is
therefore widely used in the food processing, pharmaceutical, oil drilling, biomedical, and
cosmetic industries as a biodegradable plastic, texture modifier, and thickener (Almeida
et al. 2018). This section will describe the physical characteristics of the glucomannan mol-
ecules, such as water absorption, solubility in water, viscosity, and thermal degradation
that will explain these rheological properties.
7.2.1 Water Solubility and Water Absorption

The KGM is soluble in water, but insoluble in organic solvents, such as methanol, ethanol,
acetone, or ether (Wang et al. 2015). Its solubility in water can be reduced by the formation of
strong hydrogen bonds, following purification steps or drying operations. Indeed, the most
important factor mainly affecting the aqueous solubility of the KGM is its degree of acetyla-
tion. The presence of the acetyl groups in the KGM inhibits the formation of intramolecular
hydrogen bonds, thereby improving the solubility of the KGM (Alonso-Sande et al. 2009).
This solubility may partly be attributed to the long side chains of the KGM which serve to
hinder intermolecular association and enhance solvation, but predominantly is believed
to be associated with the presence of the acetyl substituents. The precise role of the acetyl
groups in promoting solubility is still a matter of controversy (Ratcliffe et al. 2005).
The solubility can be further increased by agitating mechanically or by supplying heat
(Official Journal of the European Union 2012; Singh et al. 2018).
The konjac glucomannan exhibits a high water-absorption index: 100 g water/g sample.
It is possible to modulate such an absorption using chemical modification. In fact, as an
example of reduced hydration of the KGM in aqueous solution, acetylated konjac gluco-
mannan exhibits lower water absorption compared to the KGM (Zhang et al. 2014; Xiao
et al. 2015).
The water adsorption isotherm of the KGM sample is sigmoidal (Figure 7.3), correspond-
ing to the type II isotherms in the International Union of Pure and Applied Chemistry
(IUPAC) classification scheme (Xiao et al. 2015).
FIGURE 7.3
Experimental adsorption isotherm of the KGM at 25°C. (From Xiao, M. et al., Carbohydr. Polym., 130, 1–8, 2015.)
The interpretation of the moisture adsorption isotherms, according to the GAB model,
showed that the mono-layer moisture content in the KGM was high (12.69 ± 0.57) and the
energy constant C was 4.14 (Xiao et al. 2015).
The value of solubility and water absorption for the KGM varied due to different experi-
mental conditions and biological variation in samples (Table 7.3).
This high water-sorption capacity of the KGM leads to high viscous properties.
TABLE 7.3
Variation of Solubility and Water Absorption Values for KGM Samples
Value of Solubility for Water Absorption
Samples KGM Value References
KGM (92.6% purity, 71 ± 0.02 g dry basis in 153.64 ± 4.70 g Tatirat et al. (2012)
Mw = 1.2 × 106 Da, sample water solution/g total water/g dry basis
solution = 0.5% (w/w)) dry basis
KGM 0.829 ± 0.00185 g dry basis Chen et al. (2011)
(Mw = 7.47 × 105 g mol−1, in water solution/g total
sample solution = 0.4% sample
(w/w))
KGM (sample 105.4 g/g Koroskenyi and
solution = 0.33% (w/w)) McCarthy (2001)
KGM (sample 100 g g−1 dry sample Liu et al. (2010)
solution = 0.33% (w/w))
KGM sample 0.380 ± 0.003 g dry 38.8 ± 0.32 g Xiao, Dai et al. (2015)
solution = 0.4% (w/w)). basis/100 mL water water/g dry basis
7.2.2 Thickening Agent

In pure water, the konjac glucomannan gives highly viscous pseudoplastic solutions.
In fact, its high molecular weight and acid stability of the KGM make it an excellent
thickener for foods. It is comparable to exudate and seed gums, such as tragacanth, guar,
and locust bean gum. Because it is non-ionic, it is relatively unaffected by the levels
of salt used in many foods and is stable to below pH 3.8 (Thomas 1997). Nevertheless,
according to Wang et al. (2008), the viscosity of the KGM solution reached to the maxi-
mum when the pH value was 5.5. Moreover, the viscosity of the KGM solution (0.1% and
0.3%) decreased when the pH > 6.9. The higher the pH was, the lower the viscosity. Luo
et al. (2013) measured the intrinsic viscosity [h] and the theoretical critical concentra-
tion c* of the KGM in distilled water and found 18.91 dL/g and 0.053%wt, respectively.
They reported that in the presence of alkali, the expansion of the molecular dimen-
sion is restrained, resulting in a lower intrinsic viscosity and a higher theoretical critical
concentration.
Other authors (Jacon et al. 1993) reported that the intrinsic viscosity of the KGM was
one of the highest among the polysaccharides used in the food industries in comparison
with guar or locust bean gum solutions at the same concentration. This suggests that the
molecular mass of the konjac may be larger than that of guar and locust bean gum if their
molecules are all similar stiff chains, as suggested by light scattering studies. The role
of aggregation in enhancing viscosity, however, cannot be ruled out at the present time.
The solution viscosity can decrease with ageing, possibly by bacterial or enzymic hydro-
lysis (Nishinari et al. 1992).
According to Takigami (2009), the viscosity of 1% (w/w) KGM aqueous solution at room
temperature is 31.6 Pa.s. At the same concentration and temperature, the KGM forms
highly viscous dispersions in comparison with the other hydrocolloids for which vis-
cosities are 0.3 Pa.s for κ-carrageenan, 4.2 Pa.s for guar gum, and 8.2 Pa.s for xanthan
(Nishinari et al. 1992).
The KGM solution (1% w/v) exhibited the typical shear-thinning fluid behaviour:
decreasing apparent viscosity with shear rate (Li et al. 2017).
With the increasing shear rate, the solution KGM viscosity decreased significantly at
25°C (Figure 7.4). This shear thinning phenomenon indicated that the KGM solution is
typical pseudoplastic fluid. When concentrations of the KGM solution were below 0.55%,
the viscosity was less affected by shear rate, showing the approximate Newtonian fluid
flow characteristics (Wang et al. 2012).
After the modelling of these rheological curves with the power law (τ = K∙Dn), the values
of K and n of different concentrations of the KGM solution at 25°C are listed in Table 7.4.
When the concentration of the KGM increased, the K value increased and n value decreased.
For the KGM concentration of 0.10%, the K value was very small, the n value greater, so the
solution was very close to a Newtonian fluid (Wang et al. 2012).
Several steady shear viscosity studies were conducted on the KGM solutions of various
concentrations and showed the same results of the impact of shear rate (Dave et al. 1998;
Wang et al. 2008; Almeida et al. 2018). For all the tested KGM concentrations, a Newtonian
region, where the viscosity is independent of the shear rate, was observed at low shear
rates (Almeida et al. 2018).
The viscosity of the KGM solutions decreased for higher shear rates and displayed a
shear thinning behaviour. The viscosity of the KGM solutions increased to a great extent
with an increase in concentration from 10% to 30% (Dave et al. 1998).
FIGURE 7.4
The effect of shear rate on viscosity (η) of the KGM gum at different concentrations. (From Wang, C. et al., Phys.
Proc., 33, 25–30, 2012.)
TABLE 7.4
K and n Value of KGM Gum of Different Concentrations
Concentrations (% w/w) Viscosity Coefficient K/(mPa·s) Index n
0.10 0.006 0.9883

0.25 0.619 0.5682
0.4 3.262 0.4119
0.55 7.425 0.3640
0.70 19.343 0.3186
0.85 36.019 0.2794
1.00 58.058 0.2370
1.15 88.827 0.2314
Source: Wang, C. et al., Phys. Proc., 33, 25–30, 2012.
In fact, the viscosity of a KGM solution increases exponentially according to the polymer
concentration: the viscosity of a KGM aqueous solution at 2% (w/w) is 12 times higher than
that of a solution of the KGM at 1% (w/w) (Takigami 2009).
However, Wang et al. (2008) studied the influence of the concentration of the KGM on the
viscosity of the KGM solutions. Their results showed that the KGM concentration of 0.2%
was the threshold to observe a change of viscosity.
The viscosity of the KGM solutions is also dependent on the stirring time: two hours of
stirring are required to reach the viscosity plateau (Takigami 2009).
Other authors studied the viscosity of 30% KGM solutions as a function of temperature
for various shear rates. Quite surprisingly, it was observed that the viscosity values did
not seem to depend on processing temperatures. So, according to the authors, the KGM
solutions (concentration of 10% to 30%) can be processed indifferently at temperatures
ranging from 50°C to 90°C and (Dave et al. 1998).
FIGURE 7.5
Effect of shear rate on the viscosity of 1.00% KGM gum at different temperatures. (From Wang, C. et al., Phys.
Proc., 33, 25–30, 2012.)
The same results have been obtained by Wang et al. (2012).

The shear thinning behaviour was studied as a function of temperature (Figure 7.5).
When the temperature was lower, the shear thinning behaviour was more remarkable.
However, for a temperature higher than 85°C, the viscosity was less affected by the shear
rate, and the behaviour was close to the one of a Newtonian fluid.
Wang et al. (2008) have studied the influence of temperature from 30°C to 90°C on the
viscosity of low concentrated KGM solutions (0.1% and 0.3%). They obtained a point of
inflexion on a curve around 70°C. According to the authors, this is due to the quickened
action of the molecules with the increasing temperature, and the fact that broken mol-
ecules lose the hydrogen bonds of the KGM.
7.2.3 Thermally Stable Gels

The gelation behaviour of the KGM depends on the limited occurrence of the acetyl groups
in the copolymer. In fact, the acetyl groups inhibit the polymer chains’ aggregation and
thus slow gel formation. Upon dissolution in the alkaline coagulant, deacetylation of the
konjac flour takes place to form a thermostable gel. However, the gelation rate also depends
on the pH (9–10) and processing temperature (>200°C) (Singh et al. 2018).
During the deacetylation reaction, the storage modulus increases and the gel formation
is initiated, while the acetyl group is lost following first order kinetics. Therefore, the gela-
tion and thermal stability are highly dependent on the degree of acetylation and other
additives (Almeida et al. 2018). For example, with the addition of sodium carbonate, the
gelation time becomes shorter and the addition of sodium hydroxide promotes the gela-
tion of the KGM. Other bases for acetylation have also been studied. For example, adding
potassium carbonate to a 2% konjac solution will form a non-melting gel upon heating.
The rate of gel formation is controlled by the pH of the system and the process tempera-
ture. Usually, the pH must be raised to between 9 and 10 for gel formation. These gels
are stable to temperatures greater than 200°C (Thomas 1997). As a guideline, with the
TABLE 7.5
Relative Gel Strength Produced by Adding 10% Alkali Relative
to the Weight of Konjac in the Solution
Alkali Used % Relative Gel Strength Gel pH
Sodium carbonate 95.4 10.2

Sodium phosphate 94.9 11.4
Potassium hydroxide 90.1 12.3
Potassium carbonate 89.0 10.1
Potassium phosphate 73.3 8.1
Sodium hydroxide 31.4 12.5
Source: Thomas, W.R., Konjac gum, in A. P. Imeson (ed.), Thickening and
Gelling Agents for Food, Springer US, Boston, MA, pp. 169–179,
1997.
exception of sodium hydroxide, the best results are obtained by adding 10% alkali relative
to the weight of the konjac in the sol (Table 7.5). The pH can be lowered to 9; however, more
heat is required at a lower pH. The selection of a specific base may be strongly influenced
by the formulation of the food and the desired label. After adding the base, the mixture
must not be agitated while the gel is forming (Thomas 1997).
The gel formation of the deacetylated KGM is also dependent on the ionic environment:
the effects of cations are less significant than those of anions on gelation.
The viscoelastic behaviour of the KGM gels were studied for various KGM concentra-
tions and at 25°C. At lower frequencies, the polymer solution showed liquid-like behaviour
(G” > G’) and at higher frequencies, solid-like behaviour (G’ > G”). This is due to that
fact that at lower frequencies, the polymer chains have enough time to disentangle and
at higher frequencies, i.e., at short time periods, the polymer chains cannot disentangle
and thus form a temporary network structure. The crossover frequency (the point where
G’ = G”) is shifting to a lower frequency with the increasing KGM concentration, which is
an indication of an enhancement of that temporary network (Almeida et al. 2018).
The elastic (G’) and viscous (G”) moduli were followed as a function of the tempera-
ture for various concentrations of the KGM solution. With the increasing concentration,
both moduli increased. At a low temperature, the elastic behaviour was dominant with
G’ > G”. When the temperature increased, G” > G’ showed the disentanglement of the
KGM chains. The gelation temperature or the gel–sol transition temperature is defined
as Theat (G’ = G”), and it shifted to higher temperatures as the concentration of the KGM
increased. Almeida et al. (2018) showed that the gel–sol transition temperature, Theat
(G’ = G”), increased linearly with the KGM concentration.
It was observed that the frequency dependence of the KGM solutions decreased as the
concentration increased, especially above 7%. The system then moved from a weak gel to a
relatively strong one (Dave et al. 1998).
Yin et al. (2008) studied the effect of different sodium salts (Na2SO4, NaCl, NaNO3,
NaSCN, and Na2CO3) on the gelation behaviour of the KGM. The influence of salts on the
gelation of the KGM was quite diversiform. Even in the absence of alkali, the salting-out
sodium sulphate was found to be capable of making the KGM form a thermally irrevers-
ible gel under neutral conditions upon heating, while sodium chloride, a weak salting-out
salt, sodium nitrate and sodium thiocyanate, salting-in salts, had no such role on the gela-
tion. Upon the addition of sodium sulphate, the behaviour of the KGM in water changed
from a viscoelastic fluid at room temperature to an increasingly stiff gel at elevated tem-
peratures. The formation of the KGM gel in the presence of sodium sulphate alone may
imply a different gelation mechanism from that shown in the presence of sodium carbon-
ate via the deacetylation.
7.3 Gelling Properties with Other Polysaccharides

7.3.1 Starch-Konjac Mixed Systems
Making use of the non-reversibility property of the konjac mannan has been the basis of a
great variety of foods. The konjac mannan displays a synergistic effect with xanthan gum
to form gels when used in a ratio of 3:2, as well as with other polysaccharides, such as car-
rageenan, locust beam gum, and starch.
The only bibliographic data relating to a study between the konjac starch present in
tubers and the konjac glucomannan is presented in the work of Dorthe (2005). The author
compared the flow viscosities of the konjac flour (containing 0% to 1.1% starch) after
heat treatment by the autoclave. No difference in the viscosity value was observed.
This result assumes that no viscosity synergy exists between the konjac starch and konjac
glucomannan.
All results of the following studies were obtained from the purified konjac glucomannan.
The konjac glucomannan is characterised by a high affinity for water and a large hydro-
dynamic volume. Because of these characteristics, like most hydrocolloids, the konjac glu-
comannan cannot penetrate the starch grain. It thus remains localised in the continuous
phase which surrounds the starch granules. During gelatinization, the swelling of the
starch granules causes a reduction in the volume available for the hydrocolloid, and thus
an increase in its concentration in the continuous phase (Alloncle et al. 1989).
The effect of adding the konjac glucomannan at different levels (0.4%–1.5%) was stud-
ied during the gelatinization of various starches: wheat, maize, waxy maize, and tapioca
(7% gel). For this study: a Rapid Visco-Analyzer (Perten, Sweden) was used. For all starches,
the viscosity increased. This increase is intensified with the glucomannan concentration of
the konjac (Bahnassey and Breene 1994). Xu et al. (2012) confirmed these results by inves-
tigating the effect of the konjac glucomannan addition at 1%, 2%, and 3% on the amount of
starch in corn starch dispersion gelatinization (8%) using a rheometer. The increasing addi-
tion of the konjac glucomannan increases the viscosity peak of the matrices: a 46% increase
in the viscosity peak is observed from the addition of 1% to 2% of the konjac glucomannan
(Figure 7.6). Moreover, the gelatinization temperature is not affected. This shows that the
water retention capacity of the konjac glucomannan does not affect the initial swelling rate
of the starch granules.
Khanna and Tester (2006) studied the effect of the konjac glucomannan on the gelati-
nization of various starches (waxy maize, maize, high amylose corn, and potato) by dif-
ferential scanning calorimetry. The addition of the konjac glucomannan, regardless of the
starch tested, caused a decrease in the gelatinization enthalpy and a shift in the gelatini-
zation peak to higher temperatures. These results could be explained by an incomplete
gelatinization of the starch in the presence of the konjac glucomannan, which would
reduce the quantity of the available water necessary for the swelling of the starch gran-
ule. The same result was obtained via another method of analysis. Indeed, Funami et al.
FIGURE 7.6
Gelatinization curves of corn starch-konjac systems. The viscosity of the 8% corn starch dispersion and disper-
sions containing 1%, 2%, and 3% KGM addition were obtained using a rheometer. The solid line corresponds to
the temperature profile. (Xu, Z. et al., J. Agric. Food Chem., 60, 658–664, 2012.)
(2005) investigated the effect of the konjac glucomannan (0.5%) on the gelatinization of
wheat starch by assaying the amount of solubilised amylose in the continuous phase by
spectrophotometric assay at a wavelength of 620 nm after staining with a solution of I2 KI.
The results show that little amylose has been solubilised. The results suggest that increas-
ing the viscosity of the continuous phase prevents the diffusion of the amylose out of the
starch granule, thereby decreasing the amount of amylose released.
The influence of adding 0.5% of the konjac glucomannan to the microscopic structure of
a rice starch gel (8%) was studied by Charoenrein et al. (2011). The matrices were coloured
with a solution of fluorescein isothiocyanate dextran, and then observed by confocal laser
scanning microscopy. In a rice matrix without the konjac glucomannan, the swollen starch
grains are close, facilitating their association (Figure 7.7a). The addition of 0.5% of the kon-
jac generates starch granules less inflated, equitably distributed, and therefore a less dense
network (Figure 7.7b).
A link can be made between these microscopic observations and dynamic rheologi-
cal measurements. Yoshimura and Nishinari (1997) studied the rheological properties of
mixed corn starch and konjac glucomannan suspensions (total concentration of 3.5% poly-
saccharide with ratios of corn starch/KGM of 10/0, 9/1, 8/2, 7/3, and 6/4). They showed
that the konjac glucomannan did not interact synergistically with corn starch to promote
the formation of an ordered structure. They obtained a composite gel where the KGM is
dispersed in the continuous phase. The rheological behaviour of the KGM-corn starch
mixtures is intermediate between a concentrated polymer solution and a weak gel.
Yoshimura and Nishinari (1996) studied the physical properties of mixed corn starch and
KGM gels (total polysaccharide concentration 15%, starch/KGM ratio 13.5/1.5) by dynamic
rheology and differential enthalpic analysis. The authors showed the effect of the KGM
on syneresis fluid. The water retention capacity of the KGM prevents the occurrence of
syneresis. The maize starch retrogradation is favoured by the KGM, which absorbs water
FIGURE 7.7
Confocal electron scanning microscopy pictures of rice starch gel (8% m/m) in absence (a) and presence of 0.5%
of the konjac glucomannan (b) (scale = 20 μm). (Charoenrein, S. et al., Carbohydr. Polym., 83, 291–296, 2011.)
from the mixed gel during short-term storage (1–3 days). Beyond 7 days of storage, the ret-
rogradation of the maize starch is slowed down by the KGM. The KGM can interact with
amylose or amylopectin and prevent the formation of an ordered structure, thus delay-
ing the reorganization of amylopectin molecules. Similarly, Funami et al. (2005) observed
the retrogradation of a wheat starch-KGM mixed gel (0.5%) by differential scanning and
rheological analysis. They showed that due to the thickening properties of the KGM and
its water retention capacity, the amylose concentration in the continuous phase increases,
accelerating at ‘short-time’ the retrogradation, i.e., the amylose reorganization. On the
other hand, the KGM slows the retrogradation of starch ‘long-term’, i.e., the amylopectin
retrogradation. Xu et al. (2012) observed by differential enthalpic analysis that the KGM
delayed the retrogradation of the maize starch gels due to interactions between amylose
and the KGM.
Khanna and Tester (2006) also showed that the KGM inhibited amylopectin retrogra-
dation in starch after storage for 28 days at T < 5°C. The behaviour of the KGM is similar
to a physical barrier that prevents the association of the amylopectin chains. However,
depending on the botanical origin and the nature of the starch, the results vary. In fact,
the addition of 1% KGM almost completely inhibited the retrogradation of waxy maize
gels (0.4% amylose) and high amylose corn starch (63% amylose). In contrast, 10% of the
KGM was required to inhibit the retrogradation of corn starch gels. As for potato starch,
the retrogradation is gradually inhibited by the addition of the KGM, but to a lesser
extent.
Huang et al. (2007) studied the effect of increasing the amount of the KGM (0.1% to 0.4%)
on the rheological properties of the rice starch gels by the texture profile analysis method.
The results show no variation in the rheological parameters regardless of the concentra-
tion of the KGM. This result has been confirmed by Charoenrein et al. (2011). The authors
also showed that the addition of the KGM does not increase the firmness of the gels unlike
the addition of guar gum. This difference is due to the non-gelling property of the konjac
at low concentration or in the absence of alkaline substances.
7.3.2 Encapsulation and Complexation

Due to the large molecular weight of the KGM in the order of magnitude 105 (Kato 1969),
the KGM has a very high apparent viscosity. Besides, the KGM offers an excellent film-
forming capability, and the film formed is very stable in cold and hot water and acid solu-
tion (Dong-ying 2008). Due to this attribute, the KGM has been used as an encapsulation
material for enzymes, cells, biological agents, or mixtures thereof (Nussinovitch 2004), and
as films and membranes (Zhang 2005). However, few studies have been attempted to inves-
tigate the application of the KGM in flavour encapsulation. Yang et al. (2009) have studied
the potential application of the KGM in flavour encapsulation by taking sweet orange oil,
one of the most popular food flavours in China, as the model. These authors have shown
that because of the high apparent viscosity, the KGM must be hydrolysed before encap-
sulation and spray-drying. They highlighted that in the presence of the emulsifier Tween
80, the hydrolysed KGM offered an encapsulation yield of 82%, which is comparable with
Arabic gum and starch sodium octenyl succinate.
In the starch-KGM mixes, KGM does not form specific interactions with volatile com-
pounds. Their retention is due to the high viscosity of the continuous phase caused by the
presence of the KGM, limiting the mobility and volatility of the compounds. A salting-out
effect of the carvacrol was observed in a suspension of the KGM (Lafarge et al. 2014).
On the other hand, starch is known to develop interactions with some ligands, and this
was confirmed by Arvisenet et al. (2002), Schwartz et al. (2014), and Lafarge et al. (2017)
studies. In starchy dispersions, the intensity of retention can be explained by the ability of
each volatile compound to form specific interactions with starch, and thus depends on the
availability of amylose, but also by trapping in the complex network. Through the addition
of a small amount of the KGM to starch matrices, it might be possible to control the texture
and to extend the shelf life of food products. The addition of the KGM may also affect the
interactions between the aroma compounds and starch, and, as a consequence, affect fla-
vour retention and release.
In the work of Lafarge et al. (2014), the carvacrol retention by starch is made by the com-
plexation of amylose, whereas retention of the ethyl acetate is due to trapping by the unor-
ganised network obtained during the gelatinization. The interaction between the amylose
and ethyl acetate is mainly governed by the specific interaction with starch. The addition
of the KGM to starch dispersions is known to limit the swelling of the starch granules,
thus decreasing the solubility of the amylose. As a consequence, the retention governed by
the interaction of amylose-aroma compounds was decreased like for the carvacrol in their
study, while retention of the ethyl acetate was increased.
In a study by Schwartz et al. (2014) the effects of the KGM on the gelatinization, retrograda-
tion, and complexation properties of the potato and broad bean starches were investigated
by differential scanning calorimetry and X-ray diffraction (Figure 7.8). It was established
that the addition of 1% KGM in starch suspensions causes a slight decrease in the amount
of available water in the system. However, this seems sufficient to disrupt the gelatinization
and the retrogradation of the potato and broad bean starches. In addition, the ability of the
amylose to interact with carvacrol is strongly affected by the presence of the KGM.
The results of this study show that the presence of the KGM in a starch-water system can
affect the gelatinization of starch, the retrogradation of amylose and amylopectin, and can
disrupt the amylose complexation. This phenomenon appears essentially at low concentra-
tions of the KGM (1%), but is barely noticeable at 0.2% of the konjac. The KGM can therefore
be used to physically stabilise a starch gel with minimal disruption of the complexation
of the amylose.
FIGURE 7.8
X-ray diffraction spectra of different starch-KGM-carvacrol mixes. (From Schwartz, J.M. et al., Food Hydrocoll.,
41, 71–78, 2014.) (1) Starch 5%, (2) starch 5% carvacrol 10%, (3) starch 5% KGM 0.2% carvacrol 10%, (4) starch 5%
KGM 0.6% carvacrol 10%, and (5) starch 5% KGM 1% carvacrol 10%.
7.4 Structural Properties in Presence of Water

and Cryoprotective Properties
Normalised heating scans of the KGM-water ranging from 3% to 100% (w/w) were reported
by Dave et al. (1998). The pure KGM sample had no phase transitions from 50°C to 60°C.
Free (bulk) water exists in samples with as little as 3% of the KGM. The endotherms dis-
played upon heating were somewhat broad and structured for all the compositions, and
the transition becomes broader at higher KGM concentrations. The structure indicates that
there is water associated with the KGM. In addition, the endotherms occur partially above
0°C, suggesting different KGM-water environments, which may influence the motion and
contribute to the melting behaviour (Dave et al. 1998).
The enthalpy of melting (J g−l), ∆H, of solutions of the KGM is close to that of bulk water
(i.e., 333 J g−1) and zero at low and 100% KGM concentrations, respectively. If a linear rela-
tionship existed between the heat of fusion and % KGM concentration, then at 50% KGM
concentration, ∆H values should be about 150 J g−l. However, the experimentally determined
value was ca. 90 J g−1 (i.e., 60 J g−1 lower than expected). Therefore, the water molecules that
did not crystallise in the frozen phase (approximately 18% based on the linear relationship)
represent the non-freezing water, which is bound to the KGM backbone. This may be a rea-
son that the rheological properties were independent of the processing temperature as the
bound water molecules (18%) may act to plasticise the KGM (Dave et al. 1998).
Upon cooling, the exotherms which are attributed to the crystallization of water are
sharper at the lower KGM concentration. Significant supercooling is observed as the
freezing temperature is depressed by 15°C–20°C. The exotherms shift to lower tempera-

tures at higher concentrations, and two exotherms are observed upon cooling 40% and 50%
of the KGM concentrations. The higher and lower exotherms may be assigned to the freez-
ing of the bulk water and associated water with the KGM, respectively (Dave et al. 1998).
The KGM-water samples are not thermoreversible gels, as no melting point was observed
in the Differential Scanning Calorimetry (DSC) scans (Dave et al. 1998).
For the control of ice crystal growth, the KGM is used to replace locust bean gum, which
is widely used in dairy products, such as ice cream and cream cheese (Thomas 1997).
When rice starch gels containing 0%–0.5% KGM were subjected to five freeze–thaw cycles,
the KGM reduced the % syneresis and moderate increases in gel hardness. The scanning
electron microscopy (SEM) of the freeze–thaw gels showed the starch gel with the KGM
had smaller pores and less well defined surrounding matrices than those without the
KGM. Moreover, the confocal laser scanning microscopy images of the unfrozen gels with-
out KGM showed densely aggregated swollen starch granules, while those in gels with
KGM were more evenly distributed. These results suggest that the KGM is effective in
preserving quality in freeze–thaw rice starch gels (Charoenrein et al. 2011).
On the other hand, it has been reported that the konjac glucomannan had a cryoprotec-
tant effect on the myofibrillar protein (surimi) during frozen storage. In fact, the KGM at the
level of 1% showed the same good cryoprotective effect as a conventional cryoprotectant
(10% sucrose–sorbitol, 1:1, w/w) (Xiong et al. 2009). According to the authors, the presence
of numerous hydroxyl groups of the KGM could reduce the formation of disulphide bonds,
hydrogen bonds, and hydrophobic bonds, thus reducing the aggregation of proteins and
preventing their denaturation. In addition, the KGM, due to its good water-sorption capac-
ity, could reduce the amount of free water and therefore the amount of ice crystals formed,
thus attenuating the denaturation of proteins during storage at −18°C.
Jimenez-Colmenero et al. (2013) have studied the effect of chilled storage and freezing/
thawing on the konjac gels for use as fat replacers in the formulation of reduced/low fat
meat products. The result showed that the konjac gel showed high stability under chilling
storage conditions. The water binding capacity of the konjac ranged between 87% and 90%,
with no changes (P > 0.05) during chilling storage. The rheological behaviour and micro-
structure confirm the excellent chilling storage-stability of the KGM. In contrast to that,
the freezing/thawing process strongly affected the KGM characteristics, with a significant
decrease of the water binding properties of the KGM gels at a concentration of 5%.
After measuring the volume of the syneresis liquid, Lafarge and Journaux et al. (2017)
reported that the KGM, added in a very low quantity (0.2%) into a potato starch gel, was
able to ensure the physical stability of the gel subjected to repeated freezing/thawing
cycles. The authors explained that this result was due to the high KGM water-holding
capacity and by the competition for water between the KGM and starch.
The effect of the KGM on ice crystal growth in the KGM-based aerogels was investigated
by Ni et al. (2016). From the images obtained by low temperature polarizing microscopy
and a scanning electron microscope, the morphology and size distribution of ice crystals
were determined. The size of ice crystals was smaller and their distribution was denser
and more regular when the KGM concentration increased. In fact, without the KGM, the
aerogel structure had poor water holding capacity. Thus, the water molecules could form
ice nuclei and grow freely into large ice crystals with the migration and aggregation of
water molecules. With the addition of the KGM (0.5% (w/w)), the gel network structure
became more compact with stronger water holding capacity and may hinder water mol-
ecule migration and ice crystal growth, leading to the smallest size of ice crystals.
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8
Advances in Drying Technology
Lamul Wiset, Nattapol Poomsa-ad, Rarisara Impaprasert,

and Chaleeda Borompichaichartkul
CONTENTS
8.1 Advances in Drying Technology of Konjac Tubers....................................................... 209
8.2 Advances in Drying Technology of Konjac Flour.......................................................... 211
8.2.1 Multistage Drying.................................................................................................. 212
8.2.2 Centrifugal Technique Followed by Fluidized Bed Drying............................. 215
8.2.3 Microwave Vacuum Drying.................................................................................. 217
References...................................................................................................................................... 220
8.1 Advances in Drying Technology of Konjac Tubers

In the process of extraction of the konjac glucomannan (KGM) or purification of konjac
flour, the drying process is very important. In dry extraction, the konjac tuber is sliced into
thin chips and dried to the moisture content of 8%–10% wb (wet basis), then it is ground
into powder and sieved to remove starch and some impurities. The procedure is illustrated
in Figure 8.1.
In this step of drying, hot air or sun drying is employed to reduce the moisture content
of the wet konjac chips, while minimizing the investment and operation cost. However,
the use of some of these crude drying methods may create drawbacks with regard to the
quality of the konjac flour, such as darkening of colour, low viscosity, and non-uniform
distribution of the moisture content that could lead to the deterioration of the konjac chips
in storage. During drying, the temperature and time are the main factors affecting the
quality of the final product. Since hot air drying is one of the most frequently used meth-
ods for food dehydration, the final products are characterised by low porosity and high
apparent density (Krokida & Maroulis 1999). Moreover, hot air drying can cause heat dam-
age and significant colour changes (Krokida et al. 1998; Lin et al. 1998), as well as markedly
change the viscosity of the konjac glucomannan. Furthermore, hot air drying can take
up to several hours to dry the konjac chips. Fluidized bed drying is perhaps an alterna-
tive option to increase the drying rate of the konjac chips. Fluidization is a technique that
allows the heat and mass transfer of individual chips to occur efficiently during drying.
Montreepila (2018) studied the drying of konjac cubes using the fluidization technique.
The fluidized bed dryer that was used in the experiment is shown in Figure 8.2. It consisted
of a centrifugal fan with a three-phase electrical motor of 0.5 HP (298.3 W), with an air heating
unit installed behind the fan with six electrical heating elements (9 kW). The drying cham-
ber was an acrylic column with a 100 mm diameter. A digital temperature controller with
±0.1°C accuracy (Model YL-8N, Electric Instrument Styilong Co., Ltd, Guangdong, China)
was controlling the inlet air temperature. An inverter with ±0.1 Hz accuracy (TECO FM 5,
209
FIGURE 8.1
Dry processing of konjac flour. (From Shimizu, N., and Shimahara, H., U.S. Patent No. 3767424, 1973.)
FIGURE 8.2
Schematic diagram of the experimental fluidised bed dryer used in the experiments.
Advances in Drying Technology 211
FIGURE 8.3
Drying curves of 10 mm cubes of konjac tubers at different drying temperatures in a fluidised bed dryer of
300 mm bed thickness.
made in Taiwan) was used to control the inlet air velocity. A digital thermometer with an
accuracy of ±0.1°C (Lutron TM-903, made in Taiwan) and type K thermocouple sensor was
used.
The konjac corms were cut into 10 mm cubes using a dicer (GZZT Model MH003 China).
The initial moisture content of the sample (265.49% dry basis, db) was dried down to the
equilibrium moisture content at about 5%–6% db. The drying temperatures of 50°C, 60°C,
and 70°C with the air velocity of 2.5 m/s and the bed depth of 300 mm were the conditions
in this study. Changes in the moisture content are shown in Figure 8.3. The drying time
until the moisture content reached the equilibrium moisture content was 230, 180, and
160 minutes at the drying temperatures of 50°C, 60°C, and 70°C, respectively.
8.2 Advances in Drying Technology of Konjac Flour

The dry extraction of konjac flour has several disadvantages. One disadvantage of this
method is the difficulty to grind the dried konjac slices into a fine powder. Moreover, the
konjac flour obtained by the dry extraction method has a low extraction yield, low purity,
and low viscosity. Also, the removal of the impurities is difficult because the residues on
the surface of the corms (tachiko in Japanese) stick firmly to the konjac flour. Thus, a long
processing time is required for the comminution process (Shimizu & Shimahara 1973).
In addition, the konjac flour may be lost during the air classification process, causing a
decrease in the production yield. As a result, the konjac flour cannot be directly used as a
thickening agent and, hence, is sold as a food commodity at a low price (Chua et al. 2012).
To eliminate the drawbacks of the dry process, a wet process has been developed.
The principle of this method is to comminute raw konjac corm in a grinder in a liquid
medium, such as water or a water-miscible organic solvent, and then separate the large
particles of the konjac flour from the fine powder of tachiko by sifting (Figure 8.4). Since
the konjac flour swells and becomes viscous in a short time on contact with water, it must
be removed as fast as possible (less than 1 minute) when using water as a liquid medium in
FIGURE 8.4
Wet processing of konjac flour. (From Shimizu, N., and Shimahara, H., U.S. Patent No. 3767424, 1973.)
the grinding process. The water-miscible organic solvent may include methanol, ethanol,
propanol, acetone, and 5% of ethyl acetate-modified ethanol. N,N-dimethylformamide and
ethylene glycol dimethyl ether can also be used (Shimizu & Shimahara 1973).
The drying process for the wet extraction is important to turn the konjac slurry to pow-
der form. At this stage of drying, it is often called the secondary drying process. The com-
mon drying approach that is mostly practised, such as a hot air dryer, is inefficient and has
moderate to high energy consumption. Many researchers attempted to develop a better
drying process for drying the konjac flour for the secondary drying process of the konjac
flour. In the secondary drying process, finally, the moist konjac flour derived from the wet
extraction and purification step will undergo a drying process to become dried konjac
flour. At this stage, the improper handling of the drying process can significantly reduce
the quality of the konjac flour. The drying method, temperature, and time are the main
factors affecting the quality of the final product. Therefore, several researchers studied the
effects of drying the konjac with various degrees of success.
8.2.1 Multistage Drying

Since a material has both surface and internal moisture, the constant and falling rate periods
exist in batch drying. Therefore, the optimal drying conditions and type of dryer should
be different to remove these two distinctively different types of moisture. To remove the
surface moisture, it is generally necessary to use a more rapid process requiring a shorter
residence time in the dryer. In contrast, the internal moisture removal is a slower process,
requiring a longer residence time and often a larger capacity dryer. Normally, the first stage
may be used simply to remove the surface moisture, so that the product becomes non-sticky
and suitable for processing in the next step (Kudra & Mujumdar 2009). These considerations
are behind the concept of design and operation of a multistage drying system.
Impaprasert et al. (2013) developed a multistage drying process for producing high
quality purified konjac flour as characterised by a high whiteness index value, viscosity,
and low sulphur dioxide residue and shorter drying time. The colour of the konjac flour
changed from white to yellowish when increasing the drying time until t minutes, as
shown in Table 8.1. This fact indicates that the colour of the konjac flour will change once
the moisture ratio (Mt/Mi) was reduced to 0.4–0.6 and the temperature of the konjac flour
TABLE 8.1
Time When the Colour of Konjac Flour Changes from White to Yellowish During Hot Air Drying
Moisture Content of Konjac
Flour at Time t (min)
Time t for Colour Change of Temperature of Konjac
Drying Temp (°C) Konjac Flour (% db) Mt /Mia Flour at Time t (°C)
50 100 140.2 0.43 32.0

60 80 166.7 0.52 37.5
70 70 186.8 0.47 40.0
80 50 161.1 0.52 44.5
90 40 191.2 0.60 45.5
aMt = the moisture content on dry basis (% d.b.) at any time t during drying.
Mi = the initial moisture content (% d.b.).
TABLE 8.2
Drying Conditions in Multistage Drying Process
No. Drying Temp (°C) Drying Time for Two Stage Drying (min) Total Drying Time* (min)
1 50 300 300
2 60 280 280
3 70 210 210
4 80 145 145
5 50 + 80 100 + 110 210
6 60 + 80 80 + 95 175
7 70 + 80 70 + 70 140
* Time to reduce moisture content to 8%–10% db, the moisture ratio (Mt/Mi) = 0.03.
reached about 40°C. After this period, the colour of the konjac flour changed only slightly
during drying process. Based on this observation, the multistage can be developed by
choosing a two-stage drying process.
The drying process can be divided into two periods including before and after the colour
change. The possible drying conditions are presented in Table 8.2.
The results in Figure 8.5 show that multistage drying leads to a considerable reduction
in viscosity, with the values differing significantly (p ≤ 0.05) from the values obtained
using the conventional hot air drying at a constant drying temperature. This seems to be
due to the changes in the air temperature during the multistage drying process. A high
temperature and unsuitable drying time can cause heat damage and adversely affect the
molecular structure of the KGM. The long chain KGM is likely to be damaged by heat.
This would result in the chain length reduction and the decrease in viscosity of its solu-
tion. In comparison, freeze drying can lead to a very high viscosity (15.35 Pas). This result
shows that the drying temperature has a large effect on the quality of the konjac flour.
However, there is a trend towards increased whiteness index value with increasing the
drying temperature. When comparing between drying methods, multistage drying, con-
ventional hot air drying at a constant drying temperature, and freeze drying, the results
show that multistage drying has significantly improved the whiteness index value of the
konjac flour, whereas the whiteness index value of the freeze dried sample was very low.
This seems to be due to the shorter drying time of multistage drying. The browning of
the KGM flour is caused by exposure to the hot air. As for the residual sulphur dioxide
FIGURE 8.5
Effect of drying method and drying temperature on viscosity (a), whiteness index value (b), and sulphur diox-
ide residue content (c) of konjac flour from dried slices of A. muelleri after extraction and drying process. Letters
a, b, c on the bar graph indicate a statistically significant difference (p ≤0.05).
content, it was very low in this study and lower than the maximum limit in all samples.
In particular, the residual sulphur dioxide content in the multistage dried sample was
lower than in the conventional hot air and freeze dried samples, see results in Figure 8.5.
In addition, the maximum level of 1500 mg/kg (ppm) of sulphur dioxide is approved
by the Food and Drug Administration of Thailand for use in dried or preserved fruits
and vegetables. However, the Japanese specifications and standards for foods and food
additives indicate the maximum limit of sulphur dioxide for use with the konjac flour as
900 mg/kg (Japan External Trade Organization 2011).
8.2.2 Centrifugal Technique Followed by Fluidized Bed Drying

This work was performed by Noanlumduan (2013). The konjac corms were sliced, dried,
and ground, then extracted by the wet extraction process 3–4 times. The flour with
particle size >0.125 mm (120 mesh) was then used in the next step. The moisture in
the sample was firstly eliminated by the centrifugal technique, as shown in Figure 8.6.
The diameter of the centrifugal bucket was 30 cm with the height of 15 cm. A power
drive with a 3-phase 380-volt AC motor, 3 horsepower was used. The experiment was
conducted using the centrifugation technique at different centrifugal forces of 350, 600,
and 850 times the force of gravity.
FIGURE 8.6
Schematic diagram of centrifugal moisture extractor consisting of (1) centrifugal bucket, (2) control box, (3)
motor, and (4) set power transmissions belt.
FIGURE 8.7
Change in moisture content of konjac flour at different centrifugal forces.
The moisture reduction of the konjac flour is shown in Figure 8.7. The experimental
results from the centrifugation technique show that the centrifugal force did not affect
the moisture content of the konjac flour, in contrast to the centrifugation time that did
affect the moisture content of the konjac flour. The moisture content had rapidly decreased
within the first five minutes of centrifugation, and then remained constant. However, the
final moisture content was still high, and the material needed to be dried in a further step.
The fluidization technique is selected to be used for removing the remaining moisture
content in the konjac flour. This is possible due to the granular structure and high mois-
ture content of the sample, because this technique can remove moisture content rapidly.
The fluidised bed dryer (Figure 8.8) can rapidly reduce the moisture content, as every par-
ticle of the material is exposed to the drying air.
FIGURE 8.8
Schematic diagram of a fluidised bed dryer consisting of: (1) blower, (2) heater, (3) sieve, [120 mesh], (4) drying
chamber, and (5) sieve [120 mesh].
FIGURE 8.9
Moisture content changes of konjac flour in fluidized bed at different temperatures.
TABLE 8.3
Viscosity of Konjac Flour After Drying
Drying temperature (°C) Drying time (min) Moisture content (%wet basis) Viscosity (cP)
Sun drying – 10.36 1212.39 ± 56.52d
100 3 9.43 1455.41 ± 100.40c
120 2.5 9.09 1918.71 ± 68.51a
140 2 9.78 1810.12 ± 108.83b
Means with the different letter within a column are significantly different (p ≤ 0.05) by DMRT.
The drying temperatures of 100°C, 120°C, and 140°C were applied for the fluidization
technique. The moisture content changes of the konjac flour during drying are presented in
Figure 8.9. The experimental results using the fluidised bed dryer showed that the drying
time of 2–3 minutes reduced the moisture content of the konjac flour to less than 10% wet basis.
The viscosity of the konjac flour was assessed. The results are shown in Table 8.3.
The drying temperature significantly affects the viscosity. The higher temperature reduces
the drying time.
However, the highest temperature might not be the best condition due to the viscosity
decrease.
8.2.3 Microwave Vacuum Drying

The microwave vacuum drying is an alternative drying method, which recently became
adopted by the food industry. The heat generated by the microwave energy occurs prin-
cipally in the product, not in the oven walls or atmosphere. Therefore, the heat losses
from the oven to the surroundings are much lower, making for more comfortable work-
ing temperatures. Fast start-up and shutdown and precise process control are possible in
microwave heating (Mullin 1995; Vadivambal and Jayas 2007). The low temperature and
fast mass transfer conferred by the vacuum, combined with the rapid energy transfer by
the microwave heating, generate very rapid, low temperature drying (Yongsawatdigul &
Gunasekaran 1996; Zheng et al. 2013). Moreover, the absence of air during the drying may
inhibit oxidation, and therefore, the colour and nutrient contents of products can be largely
preserved (Nahimana & Zhang 2011; Cui et al. 2004; Kelen et al. 2006; McLoughlin et al.
2003; Sagar & Suresh 2010; Zielinska et al. 2013). Thus, a dried product with the qual-
ity equivalent to that of conventionally hot-air-dried materials can be obtained. Applying
the microwave energy under the vacuum combines the advantages of both vacuum dry-
ing and microwave drying, as far as improved energy efficiency and product quality are
concerned (Krokida & Maroulis 1999). However, most of the microwave-vacuum drying
studies focus on fruits and vegetables that need the ‘puffing’ characteristic to improve
the rehydration properties of the final product (Zhang et al. 2006). The quick microwave
energy absorption by the water molecules causes the rapid evaporation of the water from
the interior of the product towards the surface of the product, creating a flux of rapidly
escaping vapour, which helps in preventing the shrinkage and case hardening and induces
a more porous and puffing structure, thus improving the rehydration properties of the
dried materials. Markowski et al. (2009) found a higher rehydration ability for the samples
dried with microwaves under low pressure. Similar results are reported by Giri & Prasad
(2007a), who found that the rehydration properties are improved by drying at a lower
system pressure and higher microwave power, as indicated by the higher values of the
rehydration ratio. In particular, the microwave-vacuum drying techniques are reported to
be used successfully for the dehydration of many kinds of fruits and vegetables, such as
carrots (Cui et al. 2004, 2005; Nahimana & Zhang 2011), bananas (Maskan 2000; Mousa &
Farid 2002), wild cabbage (Yanyang et al. 2004), beetroot (Figiel 2010), garlic (Cui et al.
2003; Figiel 2006), mushrooms (Rodríguez et al. 2005; Giri & Prasad 2007a, 2007b), pota-
toes (Setiady et al. 2007; Song et al. 2007; Markowski et al. 2009; Song et al. 2009), mint
leaves (Therdthai & Zhou 2009), and green peas (Chauhan & Srivastava 2009; Zielinska
et al. 2013). These products possess excellent quality in terms of taste, aroma, texture, and
appearance. A number of researchers studied the effects of drying foodstuffs using this
technique with various degrees of success. The advantages of the microwave vacuum dry-
ing, especially in food and agricultural products are listed below (Kanlapong 2006):
• Efficiency: In most cases the energy couples into the solvent, not the substrate.
• Non-destructive: Drying can be done at low ambient temperatures; no need to
maintain high surface temperatures. This leads to lower thermal profiles.
• Reduction of migration: Solvents are often mobilised as a vapour, thereby are
not transporting other materials to the surface.
• Levelling effects: Coupling among wetter areas.
• Speed: Drying time can be shortened by 50% and more.
• Uniformity of drying: By a combination of more uniform thermal profiles and
levelling.
• Conveyor systems, less floor space, reduced handling: No need for batch process-
ing in most cases.
• Product improvement in some cases: Eliminates case hardening, internal stresses, etc.
As for the limitations of the microwave-vacuum drying (Kanlapong 2006), non-uniformities

in the microwave field and associated heating patterns can lead to non-uniformities in dry-
ing, which can be a significant problem, especially in regions dried earlier, non-uniformities
in the microwave field can lead to high temperatures in some regions and cause product
degradation (Lu et al. 1999). However, various ways of averaging the microwave field to
improve uniformity have been achieved by such means as mechanical movement (Torringa
et al. 1996), pneumatic agitation such as in a fluidized bed dryer (Kudra 1989), or spouted
bed dryers (Feng & Tang 1998). Some of the limitations are specific sample size and shape
(Liamkaew 2006). For industrial applications, it is difficult to dry large sizes of food and
agricultural products in the flow process because of the microwave penetration and micro-
wave leakage. The shape and size of objects heated by the microwave irradiation have much
greater and completely different impacts on the temperature distribution than the classical
means of heating. The microwave energy is produced directly in the heated material, so the
interior of the object can be heated to a higher temperature than near the surface, especially
for solids, such as frozen meat with low thermal conductivity. A number of researchers stud-
ied the effects of drying foodstuffs using this technique with various degrees of success.
In the case of the konjac flour, studies have been conducted to compare the effects of dry-
ing by various methods, such as hot air drying, multistage hot air drying, freeze drying, and
microwave-vacuum drying and related conditions on various physical quality attributes.
Those were water activity, Hunter value, whiteness index, bulk density, particle density,
porosity, apparent viscosity, glucomannan content, sulphur dioxide residues, the structure
characterization of the glucomannan flour by using the Scanning Electron Microscope
(SEM), image analyzer, and Fourier Transform Infrared (FTIR). The results from that experi-
ment indicated that the drying method had an effect on several important properties of
the konjac flour. The microwave vacuum drying seems to affect significantly physical and
structural properties of the konjac flour. This drying method decreased the bulk density
and particle density and increased the porosity of dehydrated products compared to the
conventional hot air drying. Application of the microwave vacuum drying was beneficial in
terms of reducing the processing times required and increasing the viscosity of the konjac
glucomannan solution. The highest viscosity was found after using the microwave vacuum
drying at 1440 W for 7.5 minutes. The porosity of the konjac glucomannan particles and the
viscosity of the konjac glucomannan solution seem to be related. The changes in the viscos-
ity and porosity occur in the same direction and are related to the drying conditions. It can
be observed that increasing the microwave power tends to increase the evaporation rate,
creating a flux of rapidly escaping vapour, which leads to a higher porosity. As a result, an
increase in the porosity leads the konjac glucomannan particles to absorb more water and
swell. The volume change of the konjac glucomannan particles during the water absorption
may affect the viscosity of the konjac glucomannan solution. Therefore, the more porous
granules produce more viscous solution. Similar result of the swelling in biological material
has been studied in the pharmaceutical field by Ek et al. (1995) and Hedenus et al. (2000).
They found that the porous cellulose beads were considered as a three-dimensional skeletal
fibre system on which the liquid can be taken up, both, in the pores between fibres and in the
solid fibre matrix itself. Moreover, the authors found that the pore size in the cellulose beads
almost doubled when the beads were soaked with water. For this reason, the hot-air-dried
samples with low porosity will result in a less viscous liquid. The particle shape of the konjac
flour is also an important factor affecting the viscosity of the konjac glucomannan solution.
Figure 8.10a and b shows that the microwave vacuum dried konjac flour samples have
an irregular particle shape with a rough surface. In contrast, the hot-air-dried konjac flour
sample has an oval shape with a quite smooth surface. It seems that the cavities were gener-
ated inside the particles of the konjac flour during the microwave vacuum drying process.
During the microwave vacuum drying, the heat was generated and the water was evapo-
rated to the outside of the konjac granules. Thus, the konjac granules were ruptured and
(a) (b)
FIGURE 8.10
Scanning electron micrographs of the konjac glucomannan granules after drying using a hot-air dryer at
(a) 50°C; and a microwave vacuum dryer at the power level of 1440 W (b) to the same final moisture content
5–6% db. The konjac glucomannan particle structure is shown at 350× magnification.
showed a rough shape. Hence, the ratio of the surface area to the volume of the granules
has increased and contributed to the increased viscosity of the konjac solution. In contrast,
a spherical shape possesses a minimum surface area to volume ratio resulting in reduced
cohesive forces and improved flow ability of the solution. The macrostructure observations
revealed the presence of the pores in the granules of the microwave vacuum dried kon-
jac glucomannan samples, whereas the hot-air-dried samples maintained a tightly packed
structure like in a commercial product. The colour degradation during the microwave vac-
uum drying was caused by a browning reaction. Although the microwave vacuum drying
resulted in the konjac glucomannan flour being slightly darker, the samples had a uniform
colour and no overheating or burnt spots were observed by the naked eye and no signifi-
cant difference was observed when the samples were used in the solution in food and other
applications. Given its advantages, the microwave vacuum drying has the potential for
adoption in the konjac flour industry. Using the microwave vacuum drying at a power level
of 1440 W for 7.5 minutes resulted in the best quality of the konjac flour within the range of
experimental conditions studied and provided a comparable result with the first grade of
common konjac flour in the industrial standard of China (Impaprasert et al. 2014).
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9
Konjac Industry in Major Producing Countries
Zhang Shenglin, Hadi K. Purwadaria, Chaleeda Borompichaichartkul,

CONTENTS
9.1 Konjac Industry in China.................................................................................................. 224
9.1.1 Developments in the Field Production................................................................ 224
9.1.1.1 Planting Area............................................................................................ 224
9.1.1.2 Variety Breeding...................................................................................... 224
9.1.1.3 Disease Prevention................................................................................... 226
9.1.1.4 Planting Mode.......................................................................................... 226
9.1.1.5 Planting Standardization........................................................................ 227
9.1.1.6 Major Planting Areas............................................................................... 228
9.1.1.7 Development Guidelines for Planting Industry.................................. 228
9.1.2 Current Techniques in Processing Industry....................................................... 228
9.1.2.1 Developments in Drying Techniques................................................... 228
9.1.2.2 Standards for Sulphur Dioxide Content............................................... 229
9.1.2.3 Environmental Compliance Standards................................................. 229
9.1.2.4 Impact on China of the Emergence of Konjac Production in
Southeast Asia.......................................................................................... 229
9.1.3 Development of Konjac Derived Products.......................................................... 230
9.1.3.1 Konjac Glucomannan as a Gelling Agent in the Food Industry....... 230
9.1.3.2 Product Development in Food and Other Areas................................ 230
9.1.3.3 Synchronized Expansion of Domestic and International Markets...... 230
9.1.4 The Guiding Role of National Macro-Policy....................................................... 231
9.1.4.1 Impact of Agricultural Industrialization on the Konjac Industry
in China..................................................................................................... 231
9.1.4.2 Konjac Industry Links with the National Strategy of Poverty
Alleviation................................................................................................. 231
9.1.4.3 Promotion of Konjac Products by the Healthcare Industry............... 232
9.1.5 Coordination and Management of the Industrial Associations...................... 232
9.1.5.1 Cooperation Between Key Players in the Konjac Industry................ 232
9.1.5.2 Seminars and Conferences Conducted Recently on
Konjac-Related Themes........................................................................... 232
9.1.5.3 Publicizing Konjac Industry................................................................... 233
9.1.6 Scientific Research and International Exchanges..............................................234
9.1.6.1 Scientific and Technological Achievements.........................................234
9.1.6.2 Role of International Exchanges in Konjac Industry Development..... 237
9.1.7 Market Prices........................................................................................................... 237
223
9.2 Konjac Industry in Japan................................................................................................... 238

9.2.1 Overview of the Development of Amorphophallus konjac Cultivation in Japan.... 238
9.2.1.1 Development of Konjac Planting Areas................................................ 239
9.2.1.2 Promoting Professional Production...................................................... 239
9.2.1.3 Breeding of New Varieties...................................................................... 240
9.2.1.4 Promoting Standardized Planting........................................................ 240
9.2.1.5 Improving Mechanization Level........................................................... 241
9.2.2 Overview of the Processing Industry in Japan.................................................. 241
9.2.2.1 Improve the Level of Process Automation........................................... 242
9.2.2.2 Improving the Konjac Flour Yield......................................................... 242
9.2.2.3 Separation of Foreign Materials............................................................. 243
9.2.2.4 Environmental Protection....................................................................... 243
9.2.2.5 Coordinating Role of the Japan Konjac Association........................... 243
9.2.3 Overview of the Konjac Products Industry in Japan......................................... 243
9.2.3.1 Creating Awareness about Konjac Products........................................ 244
9.2.3.2 Improving the Degree of Automation.................................................. 244
9.2.3.3 Development of New Products.............................................................. 245
9.3 Konjac Industry in Indonesia............................................................................................ 245
9.3.1 Amorphophallus Species and Their Geographic Distribution............................ 245
9.3.2 Processing Methods of Konjac.............................................................................. 246
9.3.2.1 Sun Drying of Konjac Chips................................................................... 246
9.3.2.2 Mechanical Drying of Konjac Chips..................................................... 246
9.3.2.3 Machinery for Processing of Konjac Flour........................................... 246
9.3.2.4 Production of Konjac Flour by Wet Process......................................... 247
9.3.3 Utilization of Konjac Glucomannan-Derived Products.................................... 248
9.4 Konjac Industry in Thailand............................................................................................. 250
9.4.1 Main Types of Konjac Produced and Consumed in Thailand......................... 250
9.4.2 Structure of the Konjac Industry in Thailand.................................................... 250
Acknowledgements..................................................................................................................... 252
References...................................................................................................................................... 252
9.1 Konjac Industry in China

9.1.1 Developments in the Field Production
9.1.1.1 Planting Area
At the beginning of 2008, the southern regions of China were affected by particularly cold
weather. The frost caused considerable loss of the konjac grown in Hubei, Shanxi, and
Chongqing, which reduced the konjac planting area to 1.02 million mu (15 mu = 1 ha) in
that year. The planting area returned to its original size in three years. The size of subse-
quent planting areas has expanded rapidly since then (see Table 9.1).
9.1.1.2 Variety Breeding

Over the past 10 years, China has paid particular attention to the development of the konjac
germplasm resources and made great achievements in breeding new varieties. Southwest
Konjac Industry in Major Producing Countries 225
TABLE 9.1
Planting Area of Konjac from 2008 to 2017 (unit: 10,000 mu)
Year
Type 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017
A. konjac 90 100 110 120 140 150 160 170 175 185
A. albus 12 14 15 20 30 35 40 45 50 50
Total area 102 114 125 140 170 185 200 215 225 235
University in Chongqing has published DUS Test Guide for New Plant Varieties: Konjac (dis-
tinctiveness, uniformity, and stability of certain agricultural and vegetable plants) and
established the criteria for the selection of the subsequent new plant varieties. As a result,
five new varieties of the A. konjac, one new hybrid variety, and three new varieties of the
yellow konjac were bred (see Table 9.2). The new varieties mentioned above showed great
resistance to water stress and disease and were gradually promoted and expanded in suit-
able areas.
TABLE 9.2
New Konjac Varieties Bred Between 2007 and 2017
Species/
No. Variety Name Year License No. Completed by Involved Scientists
1 A. konjac 2008 2008006 Southwest University Zhang Shenglin, Niu Yi,

Liu Haili.
2 A. konjac 2009 0003 Biotechnology & Germplasm Wang Ling, Jiang Deyou,
Resources Institute, Yunnan Li Yongjun, Ma Jiqiong,
Academy of Agricultural Sc., Qufu Yin Guifang, Chen
FuLi Developm. Co. Ltd. Jianhua
3 A. konjac 2010 2010007 Enshi Science and Technology Shian Lv, Sheng Dexian,
Bureau Liu Wenlu
4 A. konjac 2013 2013020 Chuxiong Agricultural Science Zhang Yan, Zhang
Research Institute & Suozhongzi Faxiang, Yang Faming,
Sales Agency Xu Meien
5 A. konjac 2014 2013001 Qinba Konjac Research and Cui Ming, Li Chuan,
Development Center, Ankang Wang Xianan, Zhao
University, Ankang Agricultural Xingxi, Li Zengyi
Technology Center
6 A. konjac × 2015 2015011 Enshi Agricultural Science Yang Zhaozhu, Mou
A. albus Fanggui, Liu Erxi
7 A. muelleri 2016 2016021 Kunming University, Yunnan Zhao Jianrong, Yu Lei,
Lvyuan Bio-technology Co. Ltd., Liu Yingjie
Nanjing Dianyuan Bio-technology
Co. Ltd.
7 Mi-le I 2014 2014014 Biotechnology and Germplasm Wang Ling, Sun Tao, Li
Resources Institute, Yunnan Jian, Ma Jiqiong, Chen
Academy of Agric. Yanping, Yin Guifang,
Sc.,Xishuangbanna Dai Jiang Xiaoyun, Sun
Autonomous Prefecture Institute Daowang, Yang Yi
of Agric. Sc.,Xishuangbanna Seed
Management Station
(Continued)
TABLE 9.2 (Continued)

New Konjac Varieties Bred Between 2007 and 2017
Species/
No. Variety Name Year License No. Completed by Involved Scientists
8 Mi-le II 2014 2014015 Xishuangbanna Seed Management Jian Li, Ling Wang, Tao
Station, Biotechnology and Sun, Jun Wang, Jiqiong
Germplasm Resources Institute, Ma, Shibin Wu, Fan
Yunnan Academy of Agric. Sc., Gao, etc.
Xishuangbanna Dai Autonomous
Prefecture Institute of Agric. Sc.
9 Mi-le III 2014 2014016 Xishuangbanna Dai Autonomous Tao Sun, Ling Wang
Prefecture Institute of Agric. Sc., Yanping Chem, Jian Li,
Biotechnology and Germplasm Guifang Yin, Xiaoyun
Resources Institute, Yunnan Jiang, Jiqiong Ma
Academy of Agric. Sc.,
Xishuangbanna Seed Management
Station
9.1.1.3 Disease Prevention

Since 2007, researchers in universities, research institutes, and enterprises based in China
have conducted research on konjac diseases in areas of breeding, disease identification,
disease resistance evaluation standards, biological and chemical disease control, and other
related fields. The researchers developed 13 technical standards for disease prevention, reg-
istered 24 patents for konjac cultivation and disease prevention technology, published more
than 40 papers related to konjac disease research, and obtained 46 scientific and technological
achievements awards at various levels, including 13 provincial and ministerial achievement
awards. The large-scale demonstration, promotion, and application of relevant scientific
research results have provided strong technical support for the development of China’s
konjac industry. Specific mixtures of fertilizers and anti-bacterial/anti-fungal compounds
for the konjac were developed by companies by the Hubei and Yunnan provinces. They are
now well promoted and applied in Yunnan, Sichuan, Shanxi, Hubei, Chongqing, Guizhou,
and other provinces. In addition, in the past 10 years, bacterial biocontrol agents (actinomy-
cetes D74, Streptomyces, antibiotic lysine 13-1) have been developed and are used in seed
coating and konjac-specific fertilizers. Moreover, biodiversity cultivation models, such as
the Chinese herbal medicine interplanting system, microbial fertilizers, and bio-pesticides
are being implemented as preventive measures against konjac diseases.
9.1.1.4 Planting Mode

The intercropping of the konjac and corn is the most important planting mode. It occupies
about 65% of the country’s total area. Furthermore, in the past 10 years, Ankang Shaanxi
made full use of the sparse forest resources that were returned from farmland to start
planting the konjac in the forest. Ankang achieved success with this initiative. Planting
the konjac in the forest was rapidly prevailing, and the three-dimensional planting mode
was derived in other places for konjac and other plants. The outstanding modes include
the three-dimensional planting mode of konjac from the Yichang low altitude citrus forest,
Hubei, grapes-konjac from Enshizhou, kiwi-konjac from Chongqing, Cangxi pear-konjac
from Sichuan, apple-konjac from Hanzhong, Shanxi, walnut-konjac from Yuxi, Lincang,
and Yunnan, and konjac-rubber forest from Xishuangbanna.
9.1.1.5 Planting Standardization

Standardization for konjac planting has been an important task for China for a long time.
In the past 10 years, standard planting was gradually carried out in Fuyuan and Yongshan,
Yunnan; Ankang, Shaanxi; and Jinyang, Sichuan. Fourteen technical standards were
established (see Table 9.3).
TABLE 9.3
Standards for Konjac Planting Industry
Standard
No. Standard Name Standard No. Level Completed Company
1 Local standard for DB6109/T9.(1~5)–2013 Ankang City Qinba Konjac Research and
selenium-enriched konjac Standard Development Center
cultivation soil in Ankang Ankang Agriculture Technical
Center, Ankang College
2 Determination of main DB42/T 1003–2014 Hubei Academy of Yichang Agricultural
diseases of konjac and Province Science
technical specifications Standard Academy of Enshi Agricultural
for integrated control Science
3 Technical rules for healthy DB42/T 1181–2016 Province Economic Institute of Hubei
cultivation of alpine Standard Agricultural Science, Academy of
vegetable konjac Enshi Agricultural Science,
Academy of Yichang Agricultural
Science
4 Technical rules for konjac DB42/T 1002–2014 Province Academy of Yichang Agricultural
variety test Standard Science, Academy of Enshi
Agricultural Science, Agreements
on Konjac Industry in Hubei
5 Technical rules for DB42/T841–2012 Province Hubei Jianshi Nongtai Industry Co.,
cultivation of eco Standard Ltd., Academy of Enshi Agric. Sci.,
food – konjac Academy of Yichang Agric.l Sci.,
Administration for Jianshi Science
and Technology and Administration
for Jianshi Quality Control
6 Jinyang technical rules for DB513430/T22–2014 District Jinyang A. albus Office
cultivation of konjac Standard
7 Fuyuan konjac DG5303/T5.1~8–2013 Agriculture Fuyuan Konjac Research Institute
comprehensive standard Standard in
Qujing
8 Fuyuan specification for DG5303/T5.1–2013 Agriculture Fuyuan Konjac Research Institute
cultivation of A. konjac Standard in
(seed and food corm) Qujing
9 Technical rules for storage, Q/FYQ3–2012 Company Fuyuan Jindi Konjac Seed Industry
packaging, and Standard Co., Ltd.
transportation of konjac
seed corn
10 Fuyuan technical rules for Q/FYQ2–2012 Company Fuyuan Jindi Konjac Seed Industry
konjac seed corn Standard Co., Ltd.
propagation
11 Fuyuan A. konjac corm Q/FYQ1–2012 Company Fuyuan Jindi Konjac Seed Industry
(seed and food corm) Standard Co., Ltd.
12 Technical regulations for N75–2012 District Yongshan Konjac Office and
cultivation of pollution- Standard Research Center
free A. albus in Yongshan
9.1.1.6 Major Planting Areas

In the last 10 years, the status of konjac planting in China shows a steady extension of old
plantations and also a rapid rise of new plantations. The planting industry has already
formed several traditional plantation areas in the Yunnan high-altitude area, Qinba
Mountain area, Wuling Mountain area, and Sichuan Basin surrounding mountain area.
Konjac in Anxian, Mianyang City in the Sichuan Province and Fuyuan in the Yunnan
Province won the honour of ‘Designation of Origin’. ‘Pingshan A. albus’, ‘Anxian Konjac’,
‘Emeishan Snow Konjac’, ‘Jinyang A. albus’, ‘Beichuan A. konjac’, and ‘Langao Konjac’ won
the honour of national designation of origin of agricultural products. Meanwhile, new
plantations in Huainan Guizhou, Huaihua Hunan, Luohe Henan, Shijiazhuang, and
Xingtai Hebei were gradually increased. During the past 10 years, the Konjac Association
of Chinese Society for Horticultural Science (called from here on ‘konjac association’ in this
chapter) organized three conferences to exchange expertise on konjac plantation planning
that were held in Langao (Shaanxi), Yongshan (Yunnan), and Ziyang (Shaanxi). In 2014,
Fuyuan in the Yunnan Province, Langao in the Shanxi Province, Jianshi in the Hubei
Province, Yongshan in the Yunnan Province, Zhuxi in the Hubei Province, Beichuan in the
Sichuan Province, Ziyang in the Shaanxi Province, Jinyang in the Sichuan Province, and
Liuzhi Special District in the Guizhou Province were selected as national key plantation
areas of konjac by the association.
9.1.1.7 Development Guidelines for Planting Industry

Given the low self-reproductive coefficient of the konjac and the threat of diseases to the
healthy konjac plantations, further development of the konjac industry needs to be lim-
ited until these issues have been properly addressed. Therefore, more research should
be conducted to improve the breeding of new konjac varieties, achieve a rapid propaga-
tion of improved varieties, and a better disease control. Moreover, the implementation of
standardized production methods should be promoted in konjac planting. At the same
time, with the background of obsolescence of the cultivation methods in traditional konjac
producing areas, the konjac planting industry in China needs to be modernized and a
technology transfer take place between the main producing areas. According to the local
conditions, the new plantations should be implemented by introducing mechanization
and sustainable production techniques.
9.1.2 Current Techniques in Processing Industry

9.1.2.1 Developments in Drying Techniques
At the beginning of konjac processing in China, the raw material for refined konjac flour
was produced by drying konjac slices impaled on a wooden skewer in the hot air, often
over an open fire, or sun drying konjac slices spread on the ground. With the rapidly
growing market demand for value-added processed products, the quality requirements
for konjac flour as raw material have gradually increased. The originally skewer-dried
or sun-dried konjac slices could no longer meet the market demand in terms of quality
and quantity. Therefore, the traditional drying methods had to be improved. In the last
10 years, mechanical drying systems, such as a vibrating fluidized bed dryer and mesh
belt dryer have been developed and rapidly adopted for the primary processing of konjac
in China. The use of mechanical drying equipment not only rapidly increased the quantity
of the konjac slices supplied for further processing, but also improved their quality, espe-
cially colour. This could be achieved with the help of sulphur fumigation during the dry-
ing process.
9.1.2.2 Standards for Sulphur Dioxide Content

Although the use of sulphur fumigation can reduce the browning of konjac slices dur-
ing mechanical drying, it will also leave behind sulphur dioxide residue. Therefore, the
national standard of konjac flour (GB/T18104-2000) and the agricultural industry standard
of konjac flour (NY/T 494-2002) in China set the upper limit of the sulphur dioxide residue
in the konjac flour initially at 2 g/kg. After the implementation of the new Food Safety
Law of the People’s Republic of China in 2009, it was clearly stipulated that the application
of food additives in food processing should meet the requirements of the Standard for
the Use of Food Additives (GB2760). However, the usage range of the sulphur dioxide in
processing konjac flour was not stipulated in the Standard for the Use of Food Additives
(GB2760) version 2009. Therefore, the konjac association addressed this problem jointly
with the ministry of health and the relevant agricultural department. With the concerted
efforts of all members, the Standard for the Use of Food Additives (GB2760-2014) was
issued by the Ministry of Health on the 24th of December 2014 that stipulated the sulphur
dioxide residue level permitted in the konjac flour processing. The new residue upper limit
is 0.9 g/kg. The konjac association has held several meetings to publicize the new sulphur
dioxide standard. Through the continuous research and development of equipment, the
upgrade of konjac drying equipment was completed by 2015. At the end of 2015, the asso-
ciation organized a group of experts to sample a number of konjac processing enterprises.
The results showed that the konjac flour processing from the improved equipment met the
new sulphur dioxide residue standard of 0.9 g/kg.
9.1.2.3 Environmental Compliance Standards

Attaching importance to environmental protection is the general trend of national devel-
opment. The wastes from the konjac processing must comply with related national regula-
tions. In recent years, the konjac primary drying process switched from coal combustion
to natural gas combustion. Related drying equipment has been improved and upgraded.
China also extensively promotes grinding and sifting equipment, which can effectively
remove the remaining konjac skin and buds from the refined powder. The development
and application of modern grinding processes can reduce the peeling and bud removal
procedure in the konjac washing process. Using such grinding techniques cannot only
reduce the labour cost, but also reduce the organic residue in the water after washing the
konjac. The rinsing water can be reused. After sedimentation, the waste rinsing water can
comply with the effluent standards and be used for irrigation. The dust collection equip-
ment used in konjac processing in China has also been gradually upgraded, thus reducing
the amount of dust in the processing facilities. In addition to these environmentally impor-
tant measures, noise reduction treatments will be among the key tasks on the agenda.
9.1.2.4 Impact on China of the Emergence of Konjac Production in Southeast Asia

Since the planting industry has shrunk in their home country, the Japanese business-
men started developing konjac resources in Southeast Asian countries. Soon after, some
Chinese businessmen also took an interest in the konjac resources in Southeast Asia.
They built factories to purchase and process fresh konjac in Southeast Asian countries,
such as Myanmar, Laos, Thailand, Indonesia, etc. In the past 10 years, the konjac process-
ing industry has grown rapidly in those countries. Southeast Asia became the third biggest
platform for konjac processing after China and Japan. The rise of the processing industry
in Southeast Asian countries had a marked impact on the konjac flour market in China.
On the one hand, it complements the Chinese domestic processed konjac food market, but
on the other hand, it will decrease the opportunities of exporting konjac food to Japan as
Japan will import refined konjac flour from Southeast Asian countries. Therefore, attention
should be paid to further improving the konjac processing industry in China.
9.1.3 Development of Konjac Derived Products

9.1.3.1 Konjac Glucomannan as a Gelling Agent in the Food Industry
The standards for the production of some konjac gel-based foods in China are not fully
integrated and following international standards. Therefore, there are sometimes disputes
in the domestic and international trade. The quality of the same product produced by dif-
ferent manufacturers shows significant differences. For example, the size of konjac knots,
the number of knots, the concept of the net content, and the acidity and alkalinity of pro-
tective agents do not follow a unified standard. As a result, the konjac association actively
communicates with the relevant national standard departments to implement and for-
mulate a unified standard as soon as possible. In 2009, the konjac gel food standard was
considered as a national standard project. However, the project was delayed for procedural
reasons. Later on, through the multifaceted communication of the konjac association, this
project re-attracted the relevant national departments’ attention in 2015. The project was
reviewed, and the new standard was drafted. The draft has been submitted to the relevant
national departments for reviewing. At the time of writing (2019), it is expected that the
new standard will be issued shortly.
9.1.3.2 Product Development in Food and Other Areas

The traditional konjac products in China are mainly gel-based food, including konjac tofu,
konjac shred, konjac pancake, and konjac balls. In the past 10 years, market demand has
promoted the diversification of konjac product development in China. The development of
ready-to-eat konjac food is deeply welcomed by consumers. The market is developing rap-
idly, it is even impacting the traditional spicy strip market. Since the konjac blending prod-
ucts, such as konjac granules and konjac substitute meal powder, can play a good dietary
fibre functionality role, the market demand has gradually opened up. Apart from food,
daily necessities, such as konjac cleansing cotton and konjac masks; medicines, such as
various active ingredients microencapsulated in konjac; building materials, such as konjac
putty and konjac paints, were developed over the years, showing the diversity in the use
of konjac products.
9.1.3.3 Synchronized Expansion of Domestic and International Markets

Most of the konjac products produced in China were exported to Japan over a long period
of time. From 2012 to 2013, most producing enterprises in China encountered cost increas-
ing problems, which included higher labour costs, higher packaging material prices, higher
logistics costs, and higher refined konjac flour prices. On the contrary, the sales price in the
export market did not rise concurrently. The profits were further reduced because of the
low exchange rate of the Japanese yen. On the one hand, the Chinese konjac product man-
ufacturers controlled costs by technological improvement, product innovation, and good
management, on the other hand, they paid more attention to the expansion of European,
American, Australian, and domestic markets in the crisis period. Currently, besides the
traditional exports of konjac shred to Japan, the export of konjac products, including konjac
shred and konjac pancake to Europe, America, and Australia have rapidly expanded, and
the products were well accepted. Based on the wide recognition of traditional konjac tofu
in the domestic market, people gradually understood the importance of the konjac func-
tional food rich in high-quality dietary fibre. As a result, the consumption of konjac shreds,
konjac snack foods, konjac dietary fibre granules, and other products is increasing.
9.1.4 The Guiding Role of National Macro-Policy

9.1.4.1 Impact of Agricultural Industrialization on the Konjac Industry in China
Since 2006, the Chinese government has issued a series of directives to promote the devel-
opment of agricultural industrialization in China. All regions are required to implement
the development of agricultural industrialization as an instrument of overall development
of agriculture. Konjac industrialization was put into effect in Ankang in Shaanxi; Enshi
and Yichang in Hubei; Fuyuan, Lijiang, and Chuxiong in Yunnan; Jinyang and Mianyang
in Sichuan; and Pengshui in Chongqing. This gradually led towards the advancement of
the konjac industry in China.
Agricultural industrialization has promoted the konjac industry in China by focusing
on leading industries and products, optimizing various production factors, stimulating
the regional distribution of konjac plantations, modernizing the processing industry, and
increasing the operating efficiency in the marketing of konjac products. An integration of
science, agriculture, industry, and trade had to take place involving planting, processing,
and the manufacturing industry.
Agricultural industrialization has stimulated the technological transformation of
the traditional konjac industry in China and promoted the scientific and technological
progress of the konjac industry. As a result, the konjac industry in China embarked on a
modern management mode and industrial form of self-development, self-restraint, and
self-regulation.
9.1.4.2 Konjac Industry Links with the National Strategy of Poverty Alleviation
Since 2013, poverty alleviation has become a hot social issue. The Chinese government has
promulgated a series of relevant policies to target poverty alleviation. The government has
also devised a detailed work plan focusing on poverty alleviation, which promoted the
idea of ‘Targeted poverty alleviation’. The mountainous areas inhabited by the minorities
in central and western China are characterized by relatively poor natural conditions and
a large proportion of the population with scarce economic resources. The number of eco-
nomic crops suitable for these areas is rather limited. However, konjac can be planted and
successfully grown in these areas according to the local conditions. Therefore, Yunnan,
Sichuan, Shaanxi, Hubei, Chongqing, Guizhou, Hunan, and few other provinces have
devised a series of practical measures to develop the konjac industry and promote targeted
poverty alleviation. At the same time, the recent developments in the konjac industry in
various regions have opened new perspectives for profitable konjac production within the
scope of the targeted poverty alleviation policy.
9.1.4.3 Promotion of Konjac Products by the Healthcare Industry

Healthcare plays a central role in the national development based on the current eco-
nomic development, social demand, and disease spectrum changes. It is connected with
people’s basic necessities, sickness, and death. This strategy pays attention to all kinds
of risk factors and misunderstandings that will affect health. It also advocates the com-
prehensive care of human health. It is common knowledge that as high-quality dietary
fibre, konjac glucomannan possesses important features beneficial to human health,
such as regulating the microbiological environment in the guts, promoting bowel move-
ments, and controlling weight. With this background, the konjac industry fulfils very
well its role of providing functional food with considerable benefits to the health of the
population.
9.1.5 Coordination and Management of the Industrial Associations

9.1.5.1 Cooperation Between Key Players in the Konjac Industry
In the last decade, the number of enterprises, research institutions, and self-employed
persons engaged in the development of the konjac industry increased gradually in China.
The Konjac Association of Chinese Society for Horticultural Science has created a plat-
form for all people involved in the konjac industry to exchange scientific information,
to consult technical sources, and to share know-how. The association has drawn on the
experience of the Japan Konjac Association in industrial coordination since 2008. Three
special committees were set up in the association, namely, the planting industry, pro-
cessing industry, and konjac-based product industry committees. The planting industry
committee is responsible for coordinating the location of the planting sites and following
up with the provision of agricultural extension services advising on planting technology.
They are also involved in carrying out field demonstrations and the promotion of new
varieties, new methods of disease prevention, and new planting patterns. The processing
industry committee is responsible for coordinating raw material procurement standards
for processing refined konjac flour, as well as research, development, and promotion of
new equipment and technology. This includes promoting the new standards for sulphur
dioxide content and also advising on marketing strategies by coordinating and survey-
ing the market price of konjac flour. The konjac-based product industry committee is
responsible for the development and promotion of new products and advising on pro-
duction technology and equipment, elaboration of new standards, collection and distri-
bution of market information on finished products, and forecasting the future market
demand. On this basis, the konjac association has established and improved platforms
for the communication among the commissions, expert groups, and service departments
of provinces and municipalities involved in order to strengthen the technical services
and promote information exchange. Moreover, the konjac association publishes a mem-
bers’ directory that provides useful information for all the konjac industry stakeholders
that is updated biannually. Hence, through careful division and coordination of tasks,
the overall management and service of the association are more professional, detailed,
and comprehensive.
9.1.5.2 Seminars and Conferences Conducted Recently on Konjac-Related Themes

The list of recent technical meetings related to the konjac industry is shown in Table 9.4.
TABLE 9.4
Recent Technical Meetings Related to Konjac Industry in China
Date Venue Theme
18th–20th April 2007 Chongqing Seminar on the Development of Konjac

Industry in China in 2007
3rd September 2007 Chengdu Market Analysis Meeting of Konjac Industry
17th–19th September 2008 Chongqing The Fourth Membership Conference and
Scientific & Technological Exchange
Conference
26th–28th August 2009 Chengdu (Sichuan) Seminar on the Development of Konjac
25th–28th September 2010 Chengdu (Sichuan) Advanced Training Course on Konjac
Processing Technology
5th–7th March 2010 Chongqing Principles of Konjac Cultivation, Plant
Disease Prevention, and High Yield
Technology
22nd–24th August 2010 Chongqing Seminar on the Development of Konjac
6th–8th July 2011 Langao (Shaanxi) Sixth National Konjac Planting Planning
Experience Conference
28th–29th July 2011 Yongshan (Yunnan) Seventh National Konjac Planting Planning
16th–18th September 2011 Xi’an (Shaanxi) Seminar on the Development of Konjac
21st September 2011 Beichuan (Sichuan) Coordination Meeting on Konjac Processing
25th–27th September 2012 Chongqing Fifth Membership Conference and Scientific &
Technological Exchange Conference
19th March 2013 Chengdu (Sichuan) Symposium on Implementing
GB2760 Standards for the Application &
Residue Limits of Sulfur Dioxide in Konjac
Flour Processing
12th–14th September 2013 Chongqing Symposium on The Development of Konjac
November 2013 Information not available Technical Training Course on the
Determination of Sulfur Dioxide Resides in
Konjac Flour
18th–19th September 2014 Mianyang (Sichuan) Seminar on the Development of Konjac
5th–6th August 2015 Ziyang (Shanxi) Eighth National Konjac Planting Planning
21st–22nd September 2015 Guiyang (Guizhou) Seminar on the Development of Konjac
23rd–25th September 2016 Chengdu (Sichuan) Sixth Membership Conference and
Scientific & Technological Exchange
Conference
9.1.5.3 Publicizing Konjac Industry

In order to expand the influence of the Chinese konjac industry in the whole country and
abroad, the konjac association has organized various large-scale exhibitions, such as the
China Food Exposition and Asian Food Additives Exhibition, and also held a few activities,
such as a konjac photography exhibition and annual ‘King of konjac’ selection to publicise
the konjac industry. The association calls on all members to make full use of all media to
publicise the konjac industry. In the past 10 years, the konjac industry was frequently the
object of reports and was publicised by CCTV, local TV stations, and network platforms.
On this basis, the association changed the original internal periodical ‘China Konjac
Newsletter’ into ‘China Konjac’ in 2012. The periodicals were colour printed. In order to
promote the publicity of the konjac industry in China, some information platforms were
established in the periodicals, including hot spot focus, industry dynamics, konjac cul-
ture, etc. From the 17th to 19th of July 2014, the konjac association held the first East-West
Konjac Industry Summit Forum in cooperation with local authorities in Taijiang (Guizhou
Province). On the 21st of July 2015, the association held the summit forum of the Chinese
konjac dietary fibre industry in cooperation with the China Dietary Fibre Association
and Yizhi Konjac Biotechnology Co., Ltd. On the 8th of June 2016, during the Ankang
dragon boat festival, the International Konjac Economic and Trade Fair was held by the
konjac association in collaboration with the China Association for the Promotion of
International Cooperation in Agriculture and Ankang Municipal People’s Government.
On the 17th September 2017, the association in cooperation with the Chuxiong Science
and Technology Association of Yunnan Province held the annual meeting of the konjac
industrialization and development in Chuxiong, Yunnan Province. The konjac associa-
tion has cooperated with the China Konjac Industry Network (Konjac Garden) under the
China Wangku Group. On the 21–23 September 2017, the ‘Konjac Industry Development
Summit Forum in the Internet Age’ was held with the help of the 2017 China (Sichuan)
E-commerce Development Summit. In cooperation with local governments, represen-
tatives of enterprises, and other organizations or social groups, the association will
further expand the awareness about the konjac industry by organizing large-scale con-
ferences, forums, exhibitions, and other events and also through various electronic or
printed media.
9.1.6 Scientific Research and International Exchanges

9.1.6.1 Scientific and Technological Achievements
Science and technology are the driving forces to promote the continuous development
and progress of the konjac industry in China. Over the past 10 years, China has declared
and implemented more than 200 scientific research projects about konjac at the provin-
cial and municipal levels supported by the national program ‘Factory Production and
Standardized Multiplication System Development of Konjac Test-tube Seedlings’. Nine
new varieties were bred. Four new formulations for disease prevention, such as ‘eliminat-
ing decay-konjac ling’ were developed. More than 800 patents related to konjac have been
authorized. The national standards of konjac flour and agricultural industry standards,
such as konjac flour, technical regulations for seed konjac breeding, konjac flour refiner,
and the DUS test guide for new plant varieties of konjac were elaborated. The applica-
tion for using titanium dioxide and iron oxide as colorants for konjac food processing
and sulphur dioxide as a colour protector for konjac flour processing in GB2760 was pre-
sented to the Ministry of Health. More than 900 scientific papers have been published.
Relevant scientific and technological achievements in China have been widely recognized.
Sixteen important scientific and technological awards at the provincial and ministerial
levels, including the first prize for scientific and technological progress of the Ministry of
Education, have been obtained (see Table 9.5).
TABLE 9.5
Scientific and Technological Achievements in Konjac Industry in China from 2007 to 2017
No. Achievement Name Achieved by Awarded by Year Award Category & Level
1 Research and Application of Key Technologies in Southwest University Ministry of 2007 First Prize for Scientific and
Konjac Industry in China Education Technological Progress
2 Basic Biological Research on A. konjac Southwest University Chongqing 2008 Second Prize in Natural Science
3 Study on Cultivation Techniques of Konjac with Regard Ankang Plant Protection and Ankang County 2009 Second Prize for Scientific and
to Plant Health and High Yield Quarantine Station & Southwest Technological Progress
University
4 Study on Cultivation Techniques of Konjac with Regard Qinba Konjac Research and Shanxi Province 2010 Second Prize in Science and
to Plant Health and High Yield Development Center Technology
5 Experimental and Theoretical Studies on Fine Structure Huazhong Agricultural University Hubei Province 2012 Second Prize in Natural
of Konjac Glucomannan and Its Molecular Assembly Sciences
6 Research and Application of Key Technologies in Academy of Enshi Agricultural Hubei Province 2013 Second Prize for Scientific and
Breeding and Industrialization of A. konjac Varieties in Science Technological Progress
Konjac Industry in Major Producing Countries
Qingjiang
7 Research and Promotion of Under-forest Planting Ankang Plant Protection and Shanxi Province 2015 Second Prize for Promotion of
Model of Konjac Quarantine Station Agricultural Technology
Achievement
8 Research and Promotion of Under-forest Cultivation Ankang Plant Protection and Shanxi Province 2015 Second Prize for Promotion of
Techniques of konjac quarantine Station Agricultural Technology
Achievement
9 Micro-ecological Mechanism of Healthy Growth of Northwest A&F University Shanxi Province 2017 Second Prize in Science and
Konjac Under Forest and Remediation Technology of Technology
Actinomycetes under Continuous Cropping
Conditions
10 Research and Development of Konjac Dietary Fibre Yunnan Fuyuan Jintian Agricultural Yunnan Province 2007 Third Prize for Scientific and
Products Development Co., Ltd. Technological Progress
11 A Series of Building Coatings with Konjac as Film Huazhong Agricultural University Hubei Province 2007 Third Prize for Technological
Forming Agent Invention
12 Study on Comprehensive Control Technology of Konjac Huazhong Agricultural University Hubei Province 2008 Third Prize for Scientific and
Soft Rot Technological Progress
(Continued)
235
236
TABLE 9.5 (Continued)

Scientific and Technological Achievements in Konjac Industry in China from 2007 to 2017
No. Achievement Name Achieved by Awarded by Year Award Category & Level
13 Research and Application of Integrated Prevention and Ankang Plant Protection and Society of 2009 Third Prize in Science and
Control Technology for Konjac Diseases quarantine Station Chinese Plant Technology
Protection
14 Technical Development of Konjac Dietary Fibre and Yunnan Fuyuan Jintian Agricultural Yunnan Province 2010 Third Prize for Scientific and
Glucomannan-oligosaccharide Capsule Industry Products Development Co., Ltd. Technological Progress
15 Prevention and Control of Konjac Soft Rot by Fuyuan Konjac Research Institute, Yunnan Province 2011 Third Prize for Scientific and
Biodiversity Yunnan Academy of Agricultural Technological Progress
Sciences
16 Integration and Popularization of Key Technologies for Yichang Academy of Agricultural Hubei Province 2013 Third Prize for Promotion of
Disease Resistance, High Yield, and High Quality of Sciences Scientific and Technological
A. konjac Achievements
Konjac Glucomannan
9.1.6.2 Role of International Exchanges in Konjac Industry Development

Over the past 10 years, the development of the konjac industry in China has attracted the
attention of some countries including Japan, Australia, Thailand, Myanmar, Vietnam, and
Indonesia. From the 2nd to the 6th of September 2009, the president of the konjac associa-
tion was invited to Thailand to exchange views on the development of konjac resources
and sales of konjac products in Thailand. From the 27th to the 31st of October 2009, the
president of the konjac association was invited to Myanmar to exchange views and learn
about the development of konjac resources in Myanmar. A scientist from the University
of New South Wales, Australia, involved in konjac research, visited the konjac association
twice on the 21st of December 2009 and 26th of August 2013 to have an in-depth exchange
on the development of konjac resources in China. On the 10th of June 2010, the Myanmar
visiting mission called on the association and conducted in-depth exchanges on how to
promote the development of the Myanmar konjac industry with the help of China’s expe-
rience in the konjac industry development. On the 29th of November 2012, Japanese and
Indonesian experts from the konjac community visited the association and conducted
exchanges to strengthen the cooperation between Japan, Indonesia, and China. On the
30th of July 2013, two experts representing the Japanese konjac community visited the
association. The two sides exchanged views on the development of gel food and collabo-
ration in the import and export of konjac products between Japan and China. On the 15th
of September 2014, the delegation of the Japanese konjac association from the product
branch study group visited the association to conduct in-depth exchanges between China
and Japan on cooperation in the production of konjac. From the 29th of September to the
6th of October 2015, the president of the association was invited to Vietnam to investi-
gate the konjac industry and had in-depth exchanges with the Vietnamese delegates on
the impact of the Trans-Pacific Partnership Agreement on the exchange and develop-
ment of the konjac industry between China and Vietnam. From the 9th to the 13th of
September 2016, the president of the association was invited to attend the annual meeting
of the Japanese Konjac Association and published a report, which attracted great atten-
tion of the Japanese konjac industry towards the Chinese konjac industry.
The international exchange and cooperation not only enhanced the international influ-
ence of the Chinese konjac industry, but also enabled the association to learn about the
experience of foreign countries in the development of their konjac industry, resulting in
mutual benefit to both parties.
9.1.7 Market Prices

The changes of prices of fresh tubers and dried konjac chips in China are shown in Table 9.6.
The changes of prices of konjac flour in China are shown in Table 9.7.
TABLE 9.6
Price of Fresh Konjac and Dry Konjac Chips from 2007 to 2016
Year
Type 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016
A. konjac fresh tubers (CNY/500 g) 1.1 1.3 1.5 2.0 2.2 2.2 2.1 1.8 1.3 2.2
A. konjac dry chips ten thousand CNY/tonne) 1.8 2.1 2.2 2.7 2.7 2.8 2.8 2.6 2.0 3.2
A. albus fresh tuber (CNY/500 g) 1.8 2.2 2.4 3.2 3.3 3.4 3.4 3.3 2.8 3.8
A. albus dry chips(ten thousand CNY/tonne) 2.0 2.3 2.5 3.5 3.6 3.8 3.8 4 3.5 4.2
TABLE 9.7
Price of Konjac Flour from 2007 to 2017 (Unit: 10,000 CNY/Tonne)
Year
Type 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017
A. konjac Flour 4.6 4.8 5.2 6.0 6.5 6.5 6.5 6.3 5.5 6.5 7.0
A. albus Flour 5.5 6.0 6.5 7.0 7.2 7.5 7.5 7.5 7.0 7.5 8.0
According to Tables 9.6 and 9.7, the price of the fresh tubers has doubled over the past
10 years. The price of konjac flour has increased more than 50%. Overall, the supply and
demand are both booming. In short, from 2007 to 2017 was a decade of rapid development
and growth of konjac industry. The development of the Chinese konjac industry was fruit-
ful in all respects. However, the konjac industry in China fully recognizes the bottlenecks
that still remain, hampering further development, and continue their further research and
development work to assure a healthy industry.
9.2 Konjac Industry in Japan

9.2.1 Overview of the Development of Amorphophallus konjac Cultivation in Japan
Amorphophallus konjac was introduced into Japan from China via Korea in the era of Emperor
Kimmei (539–571 AD). The coarse flour milling of A. konjac was started in the seventeenth
century in Mito (Ibaraki), with the planting of A. konjac on a large scale, and the cultivation
technology was well developed. Then large-scale cultivation of A. konjac was available in
Ibaraki, Fukuoka, Okayama, and Fukushima prefectures. Ibaraki was in first place in terms
of productivity of A. konjac in 1905. Then diseases and insect infections gave a profound
lesson to the A. konjac production. Hiroshima, Fukushima, and Okayama became the three
largest A. konjac production areas. Hiroshima took first place in 1932. And 1932 was the best
year for A. konjac production before World War II. Due to the war, the cultivation area of
A. konjac decreased down to 40% of that before the war, and gradually came back in 1957.
Gunma was the fastest growing region for A. konjac cultivation. The output of A. konjac
in Gunma achieved 46,800 tonnes, i.e., 10 times that before the war, accounting for 40% of
the national total in 1970, then 60% in 1999, and 80% after the year 2010. Concerns arose
in the Japanese A. konjac industry for the overconcentration of the plantations in Gunma.
The rapid development was attributed to the outstanding research and development, con-
tinuous breeding of new varieties, favourable natural conditions, and large pieces of lands
suitable for mechanized cultivation. In addition, due to the geographical advantages of
Gunma’s proximity to Tokyo, more support from the central government and better for-
eign trade and commercial conditions were other reasons explaining the rapid develop-
ment. According to statistics released by the Japanese Amorphophallus konjac Association,
the total cultivation area of A. konjac had increased from 11,000 hectares up to 12,200 hect-
ares, then decreased from year to year down to only 3,900 hectares in 2016. The decrease of
Amorphophallus konjac cultivation on the one hand is due to the high proportion of the age-
ing population in Japan, with the young generations not being willing to devote themselves
to farming, on the other hand, to the limited land resources. Moreover, natural disasters
like typhoons and earthquakes also have some impacts. Besides, it is also hit by the com-
petition with the steadily growing Chinese A. konjac production. Since Gunma-ken is a
prefecture with a large population and many electors, and A. konjac is the key industry of its
economy, the Japanese government set up a 1124.1% tariff to restrict the import of Chinese
Amorphophallus konjac products (http://jp.mofcom.gov.cn), and provides a government sub-
sidy to maintain the cultivation of A. konjac. As a result, the total cultivation of A. konjac in
Japan stabilized at the level of 4,000 hectares. Although the konjac industry in Japan has
declined, there are many useful and worthwhile lessons to be learned from this country.
9.2.1.1 Development of Konjac Planting Areas

In the Japanese agriculture, it was always essential to choose the appropriate location for
growing konjac. The location had to include following factors (Kurihara 1979):
• Climate: annual mean temperature of 13°C and rainfall during growing period
(May–October) between 1,000 and 1,100 mm.
• Soil: well-drained soil with rather coarse texture.
• Topography: hillside with slope in south or southeast direction at inclination of
25°–35°. The orientation and slope are related to the sunshine hours and insolation
intensity. On southwards inclined slope, the amount of solar radiation is smaller
than on the northwards inclined slope in the summer and the opposite in the
winter.
• Biological and ecological factors: trees of different height with the taller ones on
the top of the hill and shorter ones providing shading for the plantation and con-
tributing to soil conservation.
The traditional cultivation pattern is called Jinenjo (see Figure 9.1). It is a cropping pattern
resembling most the natural environment of A. konjac. Cultural practices are simple and
primitive with only mulching with wild herbs, weeding, and spraying pesticides one or
two times a year.
Among the positive characteristics of Jinenjo are the possibilities of continuous crop-
ping, low incidence of diseases, and sustainability if appropriate cultural practices are
applied. However, the hilly topography is a serious obstacle to mechanization. In spite
of the advantages of Jinenjo, Japanese agriculture is increasingly adopting the Uedama
cultivation pattern that is practiced on lowlands and allows mechanization and a more
intensive management.
9.2.1.2 Promoting Professional Production

The production unit of A. konjac in Japan and in China is a family farm. Yet, there is a huge
difference between these two countries. The major differences are as follows: first, the
Japanese farmers who grow A. konjac are very specialized, they only plant A. konjac. Second,
the Japanese plantations of A. konjac are self-sustainable. Thereby, the quality of Japanese
A. konjac is good. Third, the degree of mechanized cultivation is high, covering all agricul-
tural operations. Fourth, the supporting services are good. The agricultural associations
provide help with fertilizers and pesticides. Fifth, the sales are stable. The relationship
between the planting industry and processing plant has been established for many years.
On the contrary, in China, farmers are not specialized enough. Because of the number of
FIGURE 9.1
Schematic representation of a typical “Jinenjo” field in Japan. (Adapted from Kurihara, H., Jap. Agric. Res. Q,
13, 174–179, 1979; Chua, M., An investigation of the biology and chemistry of the Chinese medicinal plant,
Amorphophallus konjac, PhD thesis, University of Wolverhampton, Wolverhampton, UK, 2011.)
different breeding projects, the A. konjac production area is small. Consequently, the pro-
portion of mechanized cultivation is very low. The cooperatives are not popular. Most of
the sales are temporary.
9.2.1.3 Breeding of New Varieties

After World War II, Japan began to breed new varieties of A. konjac. At present, five varieties
are being grown in Japan. They include Zairai (of Japanese origin), Shina (of Chinese ori-
gin), Haruna-kuro, Akagi-ohdama, and Myogi-yutaka. The last three varieties are hybrids,
resulting from the cross-fertilization of Zairai and Shina. The Haruna-kuro and Akagi-
ohdama cultivars account for nearly 90% of the total Japanese corm production (Chua 2011).
In districts with a warmer climate, the variety Bittyu is often grown on fields with less
favourable soils. With the introduction of konjac plantations in the subtropical Okinawa
Prefecture, it is envisaged to introduce more tropical species, such as A. muelleri and A. vari-
abilis, which are known for performing better under such climatic conditions (Kurihara
1979). There are some morphological and ecological differences among ecotypes being cur-
rently grown in Japan that are taken into account in the breeding program of new varieties.
9.2.1.4 Promoting Standardized Planting

Although the natural conditions of production of A. konjac differ between China and Japan,
it is worthwhile learning from the Japanese growers about the way of standardizing the pro-
duction. First, attention needs to be paid to the quality of A. konjac seed corms; strictly do the
pre-sowing treatment, such as grading and selection in order to avoid planting damaged and
diseased corms. Second, standardize the planting: the land preparation, planting date, soil
test and fertilizer requirements, row spacing, and planting depth should be standardized.
Third, pay attention to covering seed corms well with the soil immediately after planting.
During the emergence of seedlings, open up the soil crust above the tip or use a black plastic
film covering the seedling with an opening allowing the tip to emerge. Fourth, apply disease
prevention measures. In addition to soft rot prevention, Japanese A. konjac plantations also
need protection from leaf blight caused by typhoon damage.
9.2.1.5 Improving Mechanization Level

Due to the high labour costs and high industrialization level, Japanese farmers have a high
degree of mechanized cultivation, such as grading of konjac seed corms, tillage, trenching,
planting, pesticide application, and harvesting. One person can plant more than 6 hect-
ares. China is following Japan in terms of an ageing population, rising labour costs, and
industrialization. Therefore, the mechanized cultivation of konjac in China is also the
agenda according to the actual conditions of the konjac producing areas.
9.2.2 Overview of the Processing Industry in Japan

At the beginning of the introduction of A. konjac in Japan, the tubers were directly used as
raw material to be processed into gel food. After the invention of coarse powder and fine
powder processing methods, coarse powder and fine powder became the main raw materi-
als. At the beginning of the seventeenth century, Mito (now Ibaraki Prefecture) began the
processing of konjac coarse powder. In the mid-nineteenth century, Hiroshima took the lead
in mastering the processing technology of coarse powder and fine powder, and successively
spread it to Ibaraki and Fukushima. It is noteworthy that the presentation of konjac flour pro-
cessed in Hiroshima Prefecture took place at the ‘First Domestic Trade Exposition of Japan’
held in Tokyo in 1877. The processing and sales of Japanese konjac were initially carried out
by a collaborative group of producers, which was later transformed into the Japanese Konjac
Association or Cooperative. Yet, even before the emergence of these organizations, some busi-
nessmen began to process and sell refined konjac flour. Especially in the early days of the
Showa era, the konjac industry was booming. These merchants were also raw material manu-
facturers. They not only processed the konjac raw materials in the county, but also purchased
raw materials from other regions and processed them into fine powder for sale. The trend
has gradually expanded, which has stimulated the fierce competition among raw material
processing plants. On the one hand, it has promoted the improvement of konjac processing
technology, on the other hand, the drastic changes in konjac planting have caused many raw
material manufacturers to conduct speculative trading, resulting in industrial chaos.
In 1932, in order to prevent the confusion, Hiroshima Prefecture took the lead in for-
mulating the ‘inspection rules of konjac raw materials’, and began to restrain and detect
the raw materials of konjac tubers and dried chips in market transactions. In 1940, the
Hiroshima Prefecture promulgated the testing standards of konjac flour. Since then,
Japan has unified the raw material standards of its domestic konjac industry. They have
been accepted and implemented by industry associations, governments, and enterprises.
Despite the above measures, the growth of the konjac industry has changed consider-
ably. The interests of growers, traders, and processing enterprises are diverging. At the
same time, the processing and production techniques are backward, the domestic tuber
production is insufficient, and the prices soar. Hence, for the konjac processing industry,
it is difficult to stabilize. As a result, the Japanese konjac processing industry has begun
importing raw materials from China and other countries.
Since 1912, Japanese businessmen have processed Chinese konjac raw materials into
crude powder and shipped them to Japan for further processing into fine powders.
This phenomenon is gradually prevalent in the processing and procurement circles. It has
aroused strong opposition from domestic konjac growers in Japan who appealed to the
government to prevent the import of konjac raw materials on the grounds of protecting the
domestic konjac industry. Although the Japanese government has approved the increase
of tariffs to restrict the import of konjac, the import of konjac raw materials has not been
interrupted. Once the domestic raw materials are insufficient and expensive, the proces-
sors will use the strong demand of the consumer market as the backing to coerce the
government to allow the import of raw material. Therefore, the internal contradictions
among the Japanese konjac processing cooperatives, the planting cooperatives, and the
konjac product cooperatives have not been substantially resolved.
In 1963, the Japanese Konjac Association was established. As a consortium, its subordinates
have three synergistic combinations of planting, processing, and products. It played a guid-
ing role in pre-production coordination, mid-production management, and post-production
services. Due to the shrinkage of the Japanese konjac cultivation, the number of processing
enterprises of Japanese konjac has decreased from 180 in the middle of the last century to
less than 80 at present. Fresh konjac mainly comes from Gunma and Yamanashi prefectures.
Since 1984, the production of konjac flour in Japan has experienced several twists and turns,
reaching a peak of 10,800 tonnes in 1991, and has been declining since then. By 2016, the pro-
duction was only 5,577 tonnes, which means a reduction by nearly a half.
Japanese raw material processing of konjac consisted of converting fresh konjac into
coarse powder and fine powder, which was practiced for hundreds of years. Chinese enter-
prises were processing fresh konjac into coarse powder for some time. Drawing on the
experience of Japan, in the 1980s, China developed techniques to process konjac directly
into fine powder. As a result, China has become the world’s largest producer of konjac
flour. Nevertheless, there are still various characteristics of the Japanese konjac processing
industry worth considering when designing advanced processing techniques.
9.2.2.1 Improve the Level of Process Automation

Because of the high labour cost in Japan, the konjac processing equipment has undergone
a long process of improvement by adopting advanced equipment and a high degree of
automation. All fine powder processing enterprises in Japan have achieved the one-stop
production from fresh to dried chips and to fine powder. The production line from fresh
tuber cleaning to dry chips is controlled by only one person. The slices are uniform and
the loss is minimized. The drying equipment used are efficient burners and heat exchang-
ers. The processing capacity of fresh konjac tubers is generally 1.5 tonnes/h and the yield
of fine powder is 3 tonnes/20 h.
9.2.2.2 Improving the Konjac Flour Yield

In addition to the high degree of automation, the Japanese konjac processing is also highly
sophisticated. On the one hand, it includes equipment for extracting fine powder, such as
equipment for pre-extrusion of raw material; on the other hand, it includes high-capacity
grinding and sifting equipment. The latter allows separation of fine powder and flying
powder, and thus improves the yield.
9.2.2.3 Separation of Foreign Materials

There are several foreign object inspections at various stages of the fresh tuber and fine
powder processing. This is particularly important with regard to metal detection during
fine powder processing in order to ensure the safety of the raw materials and of the pro-
cessing equipment.
9.2.2.4 Environmental Protection

Due to the lack of coal in Japan, konjac is primarily processed using heavy oil or natural
gas as fuel. This fuel has to meet stringent environmental protection requirements after
full combustion. After washing, the tubers sediments return to the field. The peeling pro-
cess produces wastes that can also be returned to the field. Because of the reasonable dust
recovery settings, the amount of dust in the factory is relatively small. The treatment of the
wastewater of the konjac gel food is strict. The wastewater is cleaned in water treatment
plants, and then recycled or used for irrigation.
Dust recovery and waste liquid standards of product processing require appropriate
equipment and investment. As for noise emissions, the processing plant should be located
far away from residential areas and necessary equipment should be added.
9.2.2.5 Coordinating Role of the Japan Konjac Association

The processing industry is in the centre of the konjac supply chain, i.e., intermediary
between the planting and the product industry. After a long-term lack of coordination,
the Japanese konjac industry gradually formed a regulatory agency, namely, the Japan
Konjac Association, which played an important role in structuring the industry. A similar
coordinating role is played by the Konjac Association of Chinese Society for Horticultural
Science (a non-profit social welfare organization).
9.2.3 Overview of the Konjac Products Industry in Japan

At first, konjac was used as medicine and snacks only among monks in Japan. It was
regarded as a precious item. With the popularization of Buddhism, it was gradually
spread from the royal family, the government, and other upper classes to the folk. After
the Edo era in Japan (1603), it became popular food for the general public. With the devel-
opment of industrial konjac processing and its products soon becoming popular, the for-
mation of the konjac industry in Japan took place earlier than that in China. After World
War II, the Japanese government attached great importance to the konjac production.
The Ministry of Health and Welfare clearly stipulated that konjac food must be added to
the meals of primary and secondary school students. Japanese konjac products are mainly
gel food. On the basis of konjac gel food, the products like konjac cake, konjac block,
konjac noodles (shirataki), konjac knots, konjac meatballs, and so on are derived. Japan
is the world’s largest consumer of konjac products thanks to the deep understanding of
the healthcare function of Amorphophallus konjac. In the Yamanashi Prefecture, there is an
exclusive shop selling konjac-based foods. More than 200 kinds of konjac foods are avail-
able that were produced by konjac processing enterprises in the Yamanashi Prefecture,
Nagano Prefecture, Kanagawa Prefecture, Gunma Prefecture, Okayama Prefecture, Aichi
Prefecture, Yamaguchi Prefecture, Gifu Prefecture, Iwate Prefecture, etc.
At present, the production of konjac in Japan mainly follows two models: production
in an automated factory and family workshop production and sales. Automated factory
production includes different steps: from the mixing and gelatinization of fine powders to
the automatic production, and the whole process of product packaging. It takes 4–5 hours to
produce different forms of konjac products, such as stripes, sheets, noodles, and meatballs.
Manufacturing enterprises require high-quality products. All products from raw mate-
rials to finished products are subject to strict quality control procedures. Factory-produced
konjac products are exported to major domestic supermarkets in Japan through the distri-
bution channels, but are also sold in exclusive shops. The production in a family workshop
is also common in the Gunma Prefecture, the main production area of konjac in Japan.
Because of the private ownership of Japanese land, several generations inherited the fam-
ily business. In the konjac industry, the façade of the family house usually hosts a char-
acteristic konjac restaurant or konjac products stores for customers to taste or purchase.
The backyard accommodates the corresponding manufacturing facilities of the konjac
products. Given its years of experience, the Japanese konjac product industry can be emu-
lated in various respects by the counterparts in other konjac producing countries.
9.2.3.1 Creating Awareness about Konjac Products

Konjac is well known in Japan. Japanese people believe that konjac is healthy. Furthermore,
Japan attaches a great importance to the konjac industrial culture. Konjac museums
have been set up in the main producing areas of Japan to host the industrial exhibitions.
The National Konjac Festival is celebrated every year on 29th May. The konjac industry is
celebrated by creating a unique atmosphere during the Konjac Festival. The promotion of
the konjac industry through large-scale activities in Japan has achieved remarkable results.
For example, at the first konjac cooking festival held in Yamagata City, on 5th September,
2010, konjac was cooked with beef and onions in a 6-meter-diameter cauldron. The dish
could feed about 30,000 people. The scene was spectacular.
In contrast to Japan, in the other producing countries including China, the creation and pro-
motion of the konjac culture is obviously insufficient. The industrial large-scale production
of konjac in China is only about 30 years old. In order to become a popular food, the general
population has to be made aware about the health benefits of konjac. For a long time, konjac
has been mainly used as a raw material to make konjac tofu to prevent starvation during wars
or famines in China. It is quite different from the current scientific understanding of konjac as
an important functional food. Therefore, at present, China and also other konjac producing
countries should make full use of multichannel advertising to increase the promotion of the
health benefits of konjac. One good way of promoting knowledge about nutritional and other
properties of konjac is by creating activities such as trade fairs and exhibitions or festivals.
9.2.3.2 Improving the Degree of Automation

Japanese labour costs are high and konjac is mainly used for gel-food production. As a
result, labour-intensive konjac products like noodles are mostly imported from China.
Therefore, Japan’s own production mainly consists of konjac-gel foods, such as konjac
blocks, cakes, or meatballs. Relying on its advanced technology, the processing equipment
in Japan is highly automated, enabling automatic control of the whole production process,
such as puffing, refining, shaping, splitting, forming, sorting, sterilization, and packag-
ing. Several studies have shown that Japan’s large-scale konjac product manufacturers can
achieve an annual production of a value exceeding 14 million USD. The factories employ
less than 20 workers, are highly automated, and have high efficiency.
9.2.3.3 Development of New Products

Japanese konjac products are relatively simple; and most of them are gel foods. Although,
in the past 10 years, konjac noodles and other products have been developed for a fast-
paced life, the function of konjac glucomannan as a valuable dietary fibre has been well
exploited. New food products, such as snack foods in small portions are being packaged
in recyclable materials that can be consumed on the run without producing waste that
would be harmful to the environment. There is also big potential for the development of
new konjac-derived non-food products for various areas, such as body care products, cos-
metics, or pharmaceuticals.
9.3 Konjac Industry in Indonesia

9.3.1 Amorphophallus Species and Their Geographic Distribution
Amorphophallus species in Indonesia have been traditionally consumed as food by low-
income communities in the majority of villages in West Java, Central Java, and East Java
in the time of food crisis, such as the occupation by foreign nations or the occurrence of
natural disasters. The Amorphophallus species grew in the wild and sufficiently for home
consumption at those various locations. Later in the 1960s, Japanese people who resided in
Indonesia and married natives started the konjac processing home industries to produce
Japanese style foods like konjac noodle and tofu. A few among well-known ones were
located in Jakarta, but later closed down. In contrast to that, in Surabaya, capital of East
Java, the production was at a larger scale, and the producers were able and continue until
now to export some of their products back to Japan.
Since the konjac-processing home industries needed a fresh konjac corm supply in
relatively large volumes, wild plants could not be relied upon anymore, and the village
communities familiar with the Amorphophallus species were encouraged to cultivate the
plants. This instigated the Amorphophallus farming, and the first major plantation, the kon-
jac agroforestry farming at the Kawasan Pemangkuan Hutan (KPH) Saradan, East Java, is
still growing larger and larger. KPH is a forest area with a defined border determined by
the Ministry of Forestry and is managed and operated under a Forest Management Unit
(FMU) set up by government.
The initial supply of fresh konjac corm was replaced by sundried konjac chips, and later
on evolved into mechanically dried konjac chips and konjac flour by the end of the 1990s,
and finally into konjac glucomannan around 2006. Konjac flour is the dried corm in pow-
der form with partially removed starch and fibre, resulting in a rather low viscosity of
the product (around 1,500 to 1,700 cps). More industries were then established at various
locations, in Jakarta, Bandung, and Surabaya, producing konjac chips, konjac flour, and
konjac glucomannan. From initially supplying domestic industries, the processed konjac
is now exported to Japan, China, and other minor importing countries, such as Korea. It is
a known fact that the konjac glucomannan industry in Indonesia uses processing machin-
ery made in China and exports the glucomannan back to China.
There are two Amorphophallus species traditionally used for food in Indonesia, i.e.,
A. variabilis and A. muelleri. Both species have been utilized for the manufacturing of
konjac noodles and tofu at the beginning of industrial era of konjac production back in
the 1950s and 1960s. A. variabilis has a white colour corm and grows more in West Java,
particularly in the Sukabumi Regency. This had been the major source of raw material for
the konjac noodle home industry in Jakarta.
A. muelleri has yellow orange colour corm and grows mostly in the East Java province
and has been so far the major source of raw material for the konjac industries of East Java.
After this, it has expanded to industries in West Java as well. This species has a higher vis-
cosity than A. variabilis. Since the late 1990s, the konjac industries in Indonesia used only
A. muelleri as a raw material for its higher viscosity characteristics.
9.3.2 Processing Methods of Konjac

9.3.2.1 Sun Drying of Konjac Chips
The conventional method of sun drying is still largely practiced among farmers under
the agroforestry scheme even though the resulting dried konjac chips show a significant
degree of dark brown colour and lower viscosity. No sodium bisulphite is applied to avoid
the darkening of the chips and the dull colour of konjac flour.
9.3.2.2 Mechanical Drying of Konjac Chips

Mechanical drying provides a better quality of konjac chips, and more so by combining
with smoking or soaking with bisulphite. The analysis of mechanically dried konjac chips
indicated that mechanical drying at 65°C for 12 hours, combined with soaking in sodium
bisulphite at 0.02% for 10 minutes, yielded yellow bright colour chips with a 12% Moisture
Content (MC) dry basis, higher glucomannan content of 55.9%, and viscosity of 7,900 cps
compared to a glucomannan content of around 20% and viscosity below 2,000 cps from
sun dried konjac chips (Purwadaria et al. 2001).
9.3.2.2.1 Tunnel Dryer

The counter-flow tunnel dryer had been designed, manufactured, and operated for a small-
scale konjac chips industry in East Java that exported the product to China. The dimension
of the tunnel dryer was 12,000 × 110 × 2,400 mm, using diesel oil as the energy source, with
a capacity of 3 ton sliced fresh konjac chips that would be dried down to 12% MC dry basis
in 12 hours at 65°C. The industry reported that the tunnel dryer increased the glucoman-
nan content from 54.0% when dried under the sun to 62.2% (Purwadaria et al. 2000).
9.3.2.2.2 Continuous Belt Dryer

A continuous belt dryer had been designed, manufactured, and operated by a medium-
scale konjac industry in East Java around 2006. The measurements of the dryer were
9,375 × 225 × 250 mm, and it had a capacity of 250 kg/h of sliced fresh konjac corm utiliz-
ing Liquefied Petroleum Gas (LPG) as the energy source. Prior to entering the dryer, the
sliced fresh konjac corm was smoked with sodium bisulphite on a belt vibrator. The indus-
try used China-made grinding equipment and exported the glucomannan back to China.
The sustainability of the industry, however, is not known.
9.3.2.3 Machinery for Processing of Konjac Flour

A series of processing machinery prototypes had been designed, manufactured, and tested
as a pilot project at the same small-scale industry where the tunnel dryer was developed.
The dried konjac chips were first fed to the rotary cutting mill, then the burr mill, the
conical ball mill, and at the end, to the screw mill. The test indicated that the flour went
through the size reduction from the volume surface diameter of 522–214 µm, and showed
an increase of the viscosity from 16,640 cps to 17,660 cps in the series of the four machines
(Purwadaria et al. 2001, 2002b).
The rotary cutting mill measuring 1,150 × 420 × 1,350 mm, powered by a 5 HP gasoline
engine as the motor, and fitted with an eccentric device to vibrate a sifter, was used to
separate starch. The capacity of the rotary mill was 65 kg dried konjac chips per hour.
The burr mill measuring 1,050 × 850 × 1,350 mm, was powered by a 5 HP gasoline
engine with 3,000 RPM motor, and had a static 70 mesh sifter. The capacity was 40 kg/h.
The conical ball mill, batch type, measuring 2,150 × 700 × 1,500 mm, and driven with
an electrical motor of 3 HP at 12 RPM. The mill was divided into three compartments.
The first and second compartment contained 25 steel balls of 60 mm diameter, and the
third compartment contained 25 steel balls of 30 mm diameter. The first compartment of
trapezoidal shape and 0.20 m length received the input material from the feeding hopper
and started the grinding. The second compartment of cylindrical shape and 0.40 m length
continued to grind the flour filling the spaces between the ball surfaces, which moved
around grinding the material. The third compartment of conical shape and 0.70 m length
finished the grinding process. The conical ball mill was also connected to a cyclone to
remove the separated starch and fibre. The capacity of the conical ball mill was 24 kg/h.
The screw mill was designed inside a horizontal tube installed on a table. It was measur-
ing at 1,250 × 520 × 1,160 mm and was driven by an electrical motor of 3 HP at 2,860 RPM.
Operating at the end of the series, the function of the screw mill was to refine the flour, and
thus it had a higher capacity than the three previous machines, namely, 665 kg/h.
A further study with a different ball mill operating at 78 RPM at a laboratory scale inves-
tigated the effects of the number and diameter of the balls and the length of milling time
(Wijanarko and Suwasito 2014). The milling process used a consecutive series of four dif-
ferent numbers and ball diameters: 21 balls of 5.4 cm, 40 balls of 4.45 cm, 78 balls of 3.5 cm,
and 245 balls of 2.4 cm. The milling time was 40, 60, 80, 100, and 120 minutes, each milling
time divided equally for the consecutive stages of different numbers and sizes of the balls.
The amount of konjac flour put into the ball mill was 1.5 kg at a ratio of 1:9 to the total
weight of the balls per one stage, namely, 13.5 kg. The results indicated that the longest
milling time had the lowest yield of 83.34% and the highest hydration capacity of 47.96%.
In the following study, Wijanarko et al. (2015) set the length of milling time to 1, 2, 3,
and 4 hours. They found that the four-hour milling time yielded the best quality of the
konjac flour that did not go through the 100 mesh sieve (70.35% glucomannan content with
16,680 cps viscosity).
In another a study, Mustafa and Wijanarko (2015) reported a higher viscosity of konjac
flour (25,410 cps) as the result of the longest milling time (L8), however, no unit of time
was provided, and the treatment was combined with an ethanol leaching process after the
konjac flour exited the ball mill. The total number of balls applied was 105 with the ratio of
4:2:1, respectively, of small-, medium-, and large-ball diameters. No specific size had been
mentioned.
9.3.2.4 Production of Konjac Flour by Wet Process

The production of konjac flour by the wet process from konjac chips has been also inves-
tigated (Purwadaria et al. 2002a). The konjac chips were ground in the rotary cutting mill,
and the powder was collected. The powder than was soaked in the water until its weight
doubled. Ethanol at various concentrations (95%, 75%, 55%, and 45% v/v) was added at vari-
ous ratios of ethanol to the water-soaked konjac powder (1/2, 1/2.5, 1/2.75, and 1/3 w/w),
and the leaching process applied two times. The slurry went through the burr mill and
the centrifuge to separate the konjac flour from the starch and fibre. The wet konjac corm
flour was dried at 70°C and processed once more in the burr mill. The results showed that
a 55% ethanol concentration with a ratio of 1/2 ethanol to the water-soaked konjac powder
resulted in the best yield among the treatments with a 67.2%–68% glucomannan content
and viscosity of 13,709 cp. The observation was done at the pilot-plant scale using the
machineries described above.
Faridah and Wijanarko (2013) optimized the leaching process using ethanol in the mac-
eration of the konjac tuber by the response surface method employing a central composite
design. Three variables were applied: leaching time, stirring speed of the homogenizer,
and the ratio of solvent as the result of extraction from the flour produced (v/w). The glu-
comannan and calcium oxalate content in the konjac flour were analyzed. The results
indicated that the response surface method predicted well the optimized condition of a
leaching time of 4 hours, 6 minutes, and 18 seconds, stirring speed of 443 RPM, and the
ratio of solvent over flour of 8.92 mL/g provided an estimate of 79.26% glucomannan and
0.07% calcium oxalate content compared to the experimental results of 79.19% and 0.08%,
respectively.
A further study (Wijanarko et al. 2014) concluded that ultrasound applied between
ethanol extractions in the following consecutive stages, 40% ethanol extraction – ultra-
sound – 60% ethanol extraction – ultrasound – 80% ethanol extraction, produced a better
quality of konjac flour. The glucomannan content, viscosity, calcium oxalate content, and
degree of whiteness obtained were at the following levels, respectively, 84.37% ± 1.79%,
13,750 ± 52.92 cps, 0.064% ± 0.004%, and 60.38 ± 0.675.
9.3.3 Utilization of Konjac Glucomannan-Derived Products

Konjac noodles and konjac tofu in the domestic industries have been produced directly
from either fresh konjac corms or dried konjac chips. In addition to that, most of dried
konjac chips, as well as konjac flour and konjac glucomannan will be exported. The food
industry in Indonesia imports back the pure glucomannan, usually from China, to be used
as ingredients in the processed products.
The utilization of konjac glucomannan-derived products, so far at the R&D stage, is
explored by a group of researchers from Brawijaya University, Malang, East Java. A pilot
plant at the Porang Research Centre, Brawijaya University, Malang, East Java has been
established, and most of the research had been carried out here. Some of the R&D publica-
tions are briefly summarized in the following paragraphs.
Wijanarko et al. (2011) analyzed the interaction of protein isolates in a food bar made
from soya, wheat, and corn protein isolates added with konjac flour for a food emergency
product by using Fourier Transform Infrared Spectroscopy (FTIR) Shimadzu 8400 S and
Scanning Electron Microscope (SEM) JSM T-100 JEOL. It was found that the food bar
together with 1% konjac flour resulted in smooth surfaces and a compact texture rather
than the rough fracture surfaces in the food bar prepared without konjac flour. Further,
the FTIR analysis indicated that interaction occurred between functional compounds of
the protein molecules in soya, wheat, and corn protein isolates and the protein molecules
of the konjac flour.
The investigation of the interaction between glucomannan and lactic acid, and between
a ƙ-casein and glucomannan-lactic acid complex, both in the water solution using molecu-
lar docking calculations, was carried out by Manab et al. (2016). The molecular docking
analysis claimed that heating at 90°C formed a complex of the glucomannan-lactic acid
and ƙ-casein with a higher binding energy compared to no heating. The lactic acid played
an important role in increasing the binding energy in its intermediation for the interaction
between glucomannan and ƙ-casein.
Several experiments of adding konjac flour into various food products have been
conducted. As a result, the recommendation was made to add konjac flour into modi-
fied cassava flour, which substituted for wheat flour as the main raw material in mak-
ing fresh noodles to improve their quality. The addition of 4% konjac flour (w/w) and
35% water (w/w) into the fresh noodle formula provided a better quality of the noodle
with 2.13 minutes cooking time, 7.03% cooking losses, 0.14 N tensile strength, 201.58%
water absorption, 103.63% volume expansion, and 51.41° of brightness (Faridah and
Wijanarko 2014).
Konjac flour of 80 mesh size produced by the wet process using ethanol as the leaching
solution was added as a mixture of ingredients with tapioca and NaCl into a formula for
meatballs. The study concluded that an ingredient composition of 5% konjac flour, 27%
tapioca, and 6% NaCl provided the best meatball physical characteristics, with 15.03 N elas-
ticity, 74.4% water holding capacity, and surface cohesiveness with small number of cavi-
ties (Dewi and Wijanarko 2015).
The addition of konjac gel as a binding agent was studied in the process of making tra-
ditional rice crackers commonly called kerupuk puli. Cooked rice was blended with other
ingredients and konjac gel, moulded to a square form, sun dried, and fried in cooking oil.
The addition of 15 g konjac corm gel with a concentration of 1% konjac flour (1 g konjac
flour per 100 mL water) into 100 g cooked rice yielded the best rice cracker of 5.77% MC
with 144.68% expansion ability during frying (Dwijanti et al. 2015).
The use of konjac corm gel at 1% concentration as a binding agent was also investigated
in the production of chicken sausage (Prastini and Wijanarko 2015). The composition of
15 g konjac corm gel with 85 g chicken meat gave the favoured chicken sausage with 98.16%
yield, 7.83 N elasticity, and 60.40% water holding capacity.
The modified konjac flour was used as an ingredient in making a kefir drink from goat
milk. The physicochemical characteristics during the storage of the kefir drink with and
without konjac flour at 4°C were investigated and compared (Manab et al. 2017). The modifi-
cation process of the konjac flour was described as follows. Three grams of konjac flour were
added to 100 mL of 8% lactic acid solution (v/v), stirred with a magnetic stirrer for 15 minutes,
irradiated in a high power microwave for 10 minutes, and then centrifuged at 5,000 RPM for
10 minutes. The results suggested that the kefir drink with modified konjac flour was more
homogenous with a smaller particle size than the kefir drink with konjac flour.
A study had been carried out using the konjac flour from yellow corm to produce restruc-
tured meat (Wijanarko et al. 2018). The konjac flour – ƙ carrageenan mixed gels – were
added to red koji rice extract in the development of the restructured meat. The optimiza-
tion of the formula was obtained using the response surface method showing that 10.21%
of the konjac flour – ƙ carrageenan mixed gels – and 6.11% red koji rice extract produced the
best restructured meat with 43.83 ± 1.82 g hardness, 72.74 ± 0.002 water holding capacity,
and 69.34 ± 0.14° hue.
The utilization of konjac glucomannan-derived products is so far still explored by the
researchers at the R&D stage. No information could be obtained whether the R&D results
have been implemented in the food processing industries. However, the Porang Research
Centre has promoted some of food processing methods using konjac flour on national TV
broadcasts, Trans 7, which can be found in the following YouTube links: https://youtu.
be/C8nXVoo3cDE for sausage processing, and https://youtu.be/xaB7h1r7rSU for jelly pro-
cessing (personal communication Wijanarko 2019).
9.4 Konjac Industry in Thailand

9.4.1 Main Types of Konjac Produced and Consumed in Thailand
At present, the konjac industry in Thailand has no central statistical records. Instead, as
far as economic data for upstream industries (tuber production), midstream industries
(processing), and downstream industries (those using konjac flour as a raw material) are
concerned, there exist estimates at the local level by the government agencies. The produc-
tion figures for all the edible Amorphophallus species growing in Thailand can be divided
into the following five categories. About 22 edible species of Konjac in Thailand can be
divided into five groups according to its application as follows:
1. The konjac which is used in starch flour industries, such as Arrowroot Starch, Buk
raw, A. saraburiensis
The konjac with tubers that can be used for making glucomannan flour apart from
A. muelleri, such as Buk Korat or Buk hu chang is A. koratensis and Buk Dang is
A. putii (in Saraburi Province) or A. linearis (in southwestern) or A. yunnanensis
(Northern). Note: A. yunnanensis is very similar in appearance to A. putii, but the
latter has a spadix subequalling the spathe and a different colour-pattern on the
petiole, peduncle, and spathe.
2. The konjac with leaf or young stem that are used for cooking, such as A. paeoniifo-
lius is mostly called elephant yam.
3. The konjac with young stem and flowers that are used for cooking, such as Buk
Teang is Arisaema petiolatum and Buk Sai Num Peung is A. elatus.
4. The konjac which is used most in the food industry for jelly manufacturing, such
as Buk Nuea Sai or Buk Khai (A. muelleri).
Buk Nuea Sai (Amorphophallus muelleri ) is a native species in Thailand, which is found in
the western and northern part of the country, such as the provinces of Kanchanaburi, Tak,
Chiang Mai, Chiang Rai, Phayao, and Mae Hong Son. This species has a high glucomannan
content (about 10% of fresh weight) and exhibits a high viscosity of its solution. Konjac glu-
comannan is in high demand from the food industries in Asian, American, and European
countries (>12,000 tonnes per year). However, Thailand can produce only about 42% of
the total demand (5,000 tonnes per year), which is not sufficient to meet the demand of
the market (Source: Kanchanaburi Agricultural Occupation Promotion and Development
Center, Highland Agricultural Extension, non-published, obtained by interview).
9.4.2 Structure of the Konjac Industry in Thailand

The commercial konjac cultivation in Thailand is concentrated in the western and north-
ern parts of the country at an altitude of 100–800 m above sea level. The konjac is planted
on the slopes or foothills in the community forest area, on well drained and well aerated
soil, and it takes more than 2–3 years until the harvest. This is why the field production
is unable to rapidly meet the increasing market demand. As a result, the farmers collect
additional konjac corms from national forest reserves. All the fresh konjac corms are pur-
chased by the processing companies for drying and milling (dry processing and produc-
tion of crude konjac flour). The pricing of the fresh konjac or dry chips is decided by the
FIGURE 9.2
Price (Baht) of fresh konjac corm per kilogram, Tak province, Thailand. (From Interview with local farmers in
Tak Province [2016].)
processors. From 2011 to 2016, the price of the fresh konjac corms has increased by about
1 Baht (1 THB = 0.03 USD) per year (Figure 9.2).
The structure of midstream konjac industries in Thailand can be divided into two cat-
egories, namely: (1) dry processing of konjac chips and (2) dry processing of glucomannan
flour. The dry processing of konjac chips in Thailand involves 2–3 major entrepreneurs
and individual SMEs with low capital, which are located in the Tak and Mae Hong Son
provinces. Most of the konjac slices products from dry processing in Thailand are pro-
cessed for export to China. In addition, many Thai farmers who have been guaranteed
the purchase price have also received from Chinese entrepreneurs the technology for the
production of konjac slices by dry processing to export to China.
In Thailand, there are few konjac glucomannan flour (KGM) processing factories
because this process requires advanced technology to produce high quality KGM flour
that is meeting the standards required by downstream industries.
As for the status of downstream industries in Thailand, most of the KGM flour is used
in the food industry, which is converting it into delicatessen products, such as noodles,
jelly, vegan food, etc. Moreover, KGM flour can be used to improve food properties, such
as gel formation in jelly and jam products, to substitute for fat in processed meat products,
to improve the texture of food, and the incorporation of KGM in dietary supplements can
be used for weight loss in a capsule form. In the biotechnology industry, konjac glucoman-
nan is used in plant tissue culture and microbial cultures. However, KGM that is used
in various industries in Thailand is mainly imported from abroad, with China being the
major source.
The overview of konjac industries in Thailand shows that the upstream, midstream, and
downstream industries are facing serious constraints because the demand from abroad is
much higher than the supply. Buk Nuea Sai (A. muelleri) is a native species in Thailand,
which is known for its high glucomannan content. Therefore, the A. muelleri corms that are
collected in the forests in addition to those harvested in the fields are not sufficient for the
market. The Thai government is currently encouraging farmers to grow more konjac in the
fields and also in the forests under the agroforestry schemes. Moreover, there is a short-
age of midstream industries in Thailand. It is found that just the dry processing of konjac
slices or processing into crude konjac flour will result in low-purity KGM. There is a need
FIGURE 9.3
Processing of konjac in Tak province, Thailand. (From Interview with local farmers in Tak Province [2016].)
for a further purification process to manufacture high-quality KGM flour. The processing
of konjac in the Tak province is shown in Figure 9.3.
In summary, it can be said that the demand for konjac flour in Thailand is rising rapidly,
but so far the supply is still low. Should, however, the upstream industry increase its out-
put of tubers by breeding high yielding cultivars, allowing plantations and agroforestry to
have enough supply, there would be a good growth potential for the midstream process-
ing industry. With a sufficient supply of raw materials, there would be higher outputs and
revenues enabling the modernization of the KGM manufacturing processes, leading to an
improvement of the quality of purified KGM. This, in turn, would benefit the downstream
sector, especially the food and pharmaceutical industry.
Acknowledgements
The authors of this chapter would like to acknowledge the valuable assistance of Dr Niu
Xinghe (COFCO Corp.), Dr Zhao Jianrong (Kunming University), and Miss Ma Jun (UNSW)
in translating the sections on the konjac industry in China and in Japan.
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10
Applications of Konjac Glucomannan
in Food and Medicine
Chaleeda Borompichaichartkul, Afwa Hayuningtyas, and Phattanit Tripetch
CONTENTS
10.1 Relevant Physicochemical Properties of KGM............................................................. 255
10.2 Utilization of KGM in Foods........................................................................................... 256
10.3 Utilization of KGM in Medicine..................................................................................... 257
10.4 Advances in Application of KGM in Food and Medicine........................................... 258
10.4.1 KGM-Milk Protein Stabilization Mechanism.................................................. 259
References...................................................................................................................................... 263
10.1 Relevant Physicochemical Properties of KGM

Konjac glucomannan (KGM) is a natural polysaccharide that is generally extracted from
konjac tubers. It has been traditionally consumed as food, but its special characteristics
allow it to be incorporated in a range of products, particularly those required in advanced
technologies. KGM solutions can have a very high viscosity depending on the glucoman-
nan content and the chain length of its molecules (Parry 2009). KGM solution is a pseu-
doplastic fluid (with viscosity decreasing as the shear rate increases) and is an excellent
thickening, gelling, and film-forming agent (Wang et al. 2012).
KGM has an excellent water solubility, yet it forms a very viscous solution at pH 5–7.
The solubility can be increased by increasing the temperature and agitation rate (European
Union 2012). However, KGM has poor solubility in organic solvents, including ethanol,
methanol, or ether (Wang et al. 2015). The viscosity of 1% (w/w) KGM solution at room
temperature is 31.6 Pa.s and increases exponentially with increasing KGM concentration.
Shah et al. (2015) reported that the viscosity of KGM solution at 2% (w/w) in water was
12 times higher than a solution with 1% (w/w) KGM. The viscosity of KGM solution was
much higher in comparison with that of other hydrocolloids at the same concentration (1%
w/w), such as guar gum was 4.2 Pa.s, κ-carrageenan was 0.3 Pa.s, and xanthan was 8.2 Pa.s
(Yoshimura et al. 1998). The reason for this phenomenon is due to the high water sorption
of KGM, namely, >100 g of water per g of KGM. The water sorption of KGM decreases with
the increase of the degree of acetylation of KGM chains (Zhang et al. 2014).
Moreover, the carbonyl and hydroxyl groups present in the molecular chain of KGM
contribute to the high solubility of KGM in water (Kohyama et al. 1993). The solubility of
a molecule in water is linked to the intermolecular hydrogen bonding, the stronger the
hydrogen bonding, the harder it is for the molecule to dissolve in water. Also, the most
255
important factor that mainly affects the solubility of KGM in water is the degree of acety-
lation. The acetyl groups potentially inhibit the intermolecular hydrogen bonds, which
improve its solubility in water (Alonso-Sande et al. 2009).
Therefore, these characteristics of the thickening and gelling agents cause konjac flour to
be popular in noodles, tofu, jelly, snack, and chewy texture foods.
10.2 Utilization of KGM in Foods

Japan and China have been using konjac tubers as food and food additives for more than
1000 years (Zhang et al. 2005). Konjac flour is used to make noodles, tofu, and traditional
snacks with springy and chewy characteristics. In Japanese cuisine, the mixture of konjac
or konnyaku flour with lime and water is used as the main ingredient of konjac Japanese
noodles (Shirataki) and other dishes, such as konjac sukiyaki and konjac udon. In tradi-
tional medicine of China, the therapeutic effects of the konjac extract are associated with
the function of detoxification, tumour-suppression, blood stasis alleviation, and phlegm
liquefaction (Niwa et al. 2000). According to Shah et al. (2015), some of the most important
KGM characteristics for human health are the ability to delay gastric emptying, aiding in
avoiding constipation, being a laxative to support diverticulitis management, being a fer-
mentable substrate in the colon, and restricting the growth of pathogens in the gut.
In the traditional way of konjac powder processing, harvested konjac tubers are washed
before peeling, then sliced, and dried by means of the sun or mechanical drying. Dried
sliced konjac are ground to produce konjac flour. Japan was the first country to develop
commercial konjac flour by pulverizing dried chips, and then using the coarse grounded
konjac chips as a raw material for traditional foods (Chua et al. 2010). The naturally odour-
less and tasteless konjac powder is easy to use in daily cooking and baking when gluten-
containing flour or other glutinous flour has to be avoided. Konjac powder can be used as
a thickening agent for sauce, glaze, soup, and stew. Nowadays, konjac flour mixed with
other gelling agents can be used as a fat analog or fat substitute. Konjac gel showed a good
ability for thermal water binding and had better elasticity during heating. It also exhib-
ited rheological properties above 40°C mimicking pork back fat (Jiménez-Colmenero et al.
2012). However, in this study, it was also observed that konjac gel presented lesser values
as compared to pork back fat of properties such as hardness, chewiness, penetration force,
gel strength, extraction force, and extrusion.
Srisamatthakarn et al. (2005) reported the use of konjac flour combined with carra-
geenan in a Makiang (fruit of Cleistocalyx operculatus) jelly recipe. The optimum combina-
tion was obtained from a Makiang fruit to water ratio of 2:1, 0.75% initial titratable acidity,
0.1% salt, and 25% sugar, with 1% konjac flour and carrageenan mixture at the ratio of 50:50.
The higher the content of the konjac-carrageenan mixture, the higher the gel strength of
the jelly. No changes were observed among other physical and chemical quality attributes
of the Makiang jelly. Akesowan and Choonhahirun (2014) studied the optimization of the
ratio of konjac gel and κ-carrageenan and sweeteners in orange jelly. They found that the
optimal conditions for the highest gel strength were 1% konjac/κ-carrageenan (39.56:60.44)
with 9.58% sucrose, 1% konjac/κ-carrageenan (39.87:60.13) with 2% xylitol, and 1% konjac/κ-
carrageenan (38.18:61.82) with 0.94% erythritol-sucralose. In addition, the orange konjac jel-
lies produced under any of the three optimal conditions showed no significant differences
in appearance and colour. In a similar study, Phoonphun (2004) reported that the mix of
carrageenan and glucomannan in the ratio of 9:1 at 0.20% in the strawberry flavoured milk
Applications of Konjac Glucomannan in Food and Medicine 257
jelly drink (skim milk: strawberry juice, 55:45) produced the highest textural scores (hard-
ness 036 N., 0.81 N.mm of internal consistencies, and 8.97% of syneresis) and the highest
overall liking score. Therefore, konjac glucomannan produces better results in combina-
tion with carrageenan when trying to obtain a textural property that will be accepted by
consumers without interference in taste or appearance of the product.
The work of Impaprasert et al. (2017) studied the effects of alkalinity using limewater
versus calcium hydroxide and the gelling agent sodium alginate on the textural proper-
ties of konjac noodles. Drying and rehydration conditions were studied to evaluate the
optimum conditions for producing dried konjac noodles. By considering the springiness
and cohesiveness of the konjac noodles, their results indicated that using 3% konjac gluco-
mannan flour with limewater and an incubation time of 30 minutes were the most suitable
conditions. In addition, hot air drying at 80°C for 55 minutes and soaking in hot water for
9 minutes were the optimum drying and rehydration conditions.
The application of KGM in food is favoured by many food manufacturers since KGM
is tasteless, odourless, and colourless. The European Union permits the use of KGM as
food additive no. E-425 as a thickening agent, gelling agent, emulsifier, stabilizer, and film-
forming agent (Behera & Ray 2016). In the pharmaceutical industry, KGM can be used
for drug delivery, cellular therapy, prosthetic implants, cosmetics, and sound absorption
(Zhang et al. 2005).
KGM has attracted more attention as a dietary fibre due to its non-harmful and non-toxic
properties, good biocompatibility, functional properties, and health benefits (Behera &
Ray 2016). Since KGM shows the most positive effects, it is a potential fat replacer for use
in mozzarella cheese in terms of functional properties (Dai et al. 2018). Furthermore, Dai
et al. (2019) reported that KGM may be used as a fat replacer in manufacturing fat-reduced
mozzarella cheeses to improve functionality and pizza bake characteristics. The authors
studied the effects of KGM addition on functional and pizza bake properties of low-fat
and skimmed mozzarella cheese during refrigerated storage. KGM addition in low-fat and
skimmed mozzarella cheeses decreased the firmness, but did not affect the stickiness of
the cheese blocks. Free oil formation was not changed by KGM addition. The cheeses with
KGM addition showed more desirable pizza bake performance, as they exhibited more
complete shred melt and less scorching on the cheese surface than the cheeses without
KGM.
Recently, a degraded product of KGM, depolymerized KGM, has attracted attention
because of its low viscosity, improved hydrophilic properties, and favourable physiologi-
cal functions. Depolymerized KGM has been widely studied for health benefits, such as
the effect of a prebiotic and other relevant factors. Potential applications of depolymer-
ized KGM in the fields of anti-oxidation and immune function are also considered (Jiang
et al. 2018)
10.3 Utilization of KGM in Medicine

KGM has been used for a long time in some parts of Asia as food and also as medicine.
In traditional Chinese medicine, the KGM gel has been used for detoxification, tumour
treatment, and decreasing blood pressure, and for decades it has been used in China to
treat several diseases, such as cough, asthma, breast pain, skin disease, and hernia (Chua
et al. 2010). The introduction of KGM as a food additive and a small-scale supplement
to the United States and Europe was established in the past two decades, followed by
TABLE 10.1
Application of KGM in Food and Medicine
Function
Thickening Agent/ Cholesterol
Product Gelling Agent Texture Fibre Weight Control Blocking
Noodles/pasta ✓ ✓ ✓ ✓
Meat ✓ ✓
Bread/biscuit ✓ ✓ ✓ ✓ ✓
Ice cream ✓
Jam/marmalade ✓
Drink (fruit juice) ✓ ✓
Sauce/gravy ✓ ✓
Cup jelly ✓ ✓
Pudding/mousse ✓ ✓
Capsule, pill, tablet, ✓ ✓ ✓
or liquid
Source: Tester, R. and Al-Ghazzewi, F., Food Hydrocoll., 68, 246–254, 2017.
formulations in the form of capsules, tablets, and beverages. Moreover, initial clinical
studies have established that introducing KGM to the diet can significantly contribute
in decreasing plasma cholesterol, enhancing carbohydrate metabolism, colonic ecology,
and bowel movements (Chua et al. 2010). As a food additive, KGM is authorized in
Europe as E-425. KGM is declared as ‘generally recognised as safe’ by the Food and Drug
Administration (FDA). It also presents excellent functional properties and is claimed as
a low-calorie ingredient due to its non-digestible fibre compound. Therefore, KGM has a
great potential to be developed as a functional food with various health benefits (Jiménez-
Colmenero et al. 2012). KGM is also reported as a largely used emulsifier and stabilizer in
food, drinks, and cosmetic products due to its gelling properties and particular rheologi-
cal properties (Behera & Ray 2017). Moreover, the suitability of KGM to be used for drug
delivery has also been thoroughly studied, especially with regard to controlled release
properties, bioadhesive properties, and cellular therapy (Zhang et al. 2014). Table 10.1 sum-
marizes the different applications of KGM in food and food supplements.
10.4 Advances in Application of KGM in Food and Medicine

Originally, the purified konjac flour was used in food production, and now it is also used
as a food additive, especially as a gelling and thickening agent that is permitted as a food
ingredient in Europe with the E-425 number (FDA 2016). The special rheological and gell-
ing properties of KGM were suitable for its use as an emulsifier and stabilizer in food, bev-
erages, cosmetics, and pharmaceuticals. Since 1996, KGM was also approved as a binder
in meat and poultry products by the United States Department of Agriculture (USDA).
KGM and its derivatives have been broadly exploited in the pharmaceutical industry for
drug delivery with its controlled release ability (Wu & Shen 2001). In the biotechnology
area, KGM is applied as a multipurpose edible film, which promotes the concept of the
probiotic edible film. As knowledge is growing, KGM can also be used in advanced tech-
nology, such as encapsulation, microencapsulation, and nanoencapsulation of bioactive
materials. Nussinovitch (2004) invented the hydrocolloid membrane with KGM in its for-
mulation. The thermally stable droplet was able to encapsulate a liquid containing at least
one enzyme, a cell, a biological agent, a pharmaceutical component, an immunological
agent, or the mixture of those compounds. This hydrocolloid membrane was able to hold
the liquid containing essential compounds without bursting at temperatures −20°C–90°C.
The concept of encapsulation was further observed at a micro scale. The aim of recent
research on KGM is to use it as a coating material in microencapsulation via the spray dry-
ing. Microencapsulation is a technique where small particles or droplets are surrounded
by a wall material or embedded in the homogeneous or heterogeneous matrix and provide
small capsules with many useful characteristics. In food technology, microencapsulation
can be used for liquid, solid, and gas droplets that are entrapped into thin films of a food
grade microencapsulating agent (Gharsallaoui et al. 2007). Konjac glucomannan is a poten-
tial agent to be used as wall material. The mechanism will be explained in Chapter 11.
10.4.1 KGM-Milk Protein Stabilization Mechanism

KGM is classified as a poor surface-active polysaccharide. In many emulsion systems,
the formation, functional properties, and stability of the emulsion can be improved using
combinations of emulsifiers (Lafarge & Cayot 2018). Previous studies have shown that
introducing polysaccharides into the water phase of protein stabilized emulsion can
improve the stability of the emulsion (Dickinson 2011). The system with the combina-
tion of emulsifiers can give the maximum advantage of the proteins, which are more
surface-active in the emulsion formation, and polysaccharides, which can stabilize the
emulsion by steric repulsion or viscosity enhancement. In fact, both proteins and poly-
saccharides contribute to enhancing the properties of emulsion and improving the sta-
bility and functionality by their emulsifying and stabilizing abilities (Bouyer et al. 2012,
Lafarge & Cayot 2018).
Figure 10.1 shows the KGM-milk protein stabilization mechanism.
The stabilization mechanism of polysaccharide-protein stabilized emulsion is mostly
due to complex formation by either covalent bonding or electrostatic interaction (Bouyer
et al. 2012). However, in KGM-protein stabilized emulsion, the stabilization mechanism
can be by polymeric stabilization (without complexation) since KGM is a non-ionic
FIGURE 10.1
Illustration of protein-stabilized emulsion with KGM. (Lu, W. et al., Food Hydrocoll., 81, 120–128, 2018.)
polysaccharide. The stabilization mechanism of KGM-protein stabilized emulsion consists

of two-step mechanisms (Bouyer et al. 2012; Hu et al. 2016).
1. Protein adsorption: The protein allows the formation of small droplets during the
emulsification process. The hydrophobic part of the protein is denatured at the oil-
water interface, which is followed by fixing on the surface of the oil in the emulsion
2. Polysaccharide stabilization (KGM): The hydrophobic part will interact with the
adsorbed protein by hydrophobic interaction, and the predominant hydrophilic
part will be oriented to the water phase, which generates an extended thickening
network, which induces a high viscosity at a low shear rate, which then enhances
the steric hindrance and force of friction between droplets and the continuous
phase, thus slowing down the droplet motion of the system.
By this mechanism, both proteins and polysaccharides contribute through their

emulsifying and stabilizing properties to enhance the stability and functionality of
the emulsions. The amphiphilic mixture increases the viscosity of the water phase and
is adsorbed well at the oil-in-water interface with the hydrophobic side chains oriented
to the oil phase and the hydrophilic side chains oriented to the water phase (Lafarge &
Cayot 2018). These properties indicate that by introducing KGM to the water phase of
emulsions, this could make it possible to design curcumin emulsion in milk systems con-
taining medium chain triglyceride (MCT) oil with potential emulsion stability and control
release properties for use in medicinal purposes. Hayuningtyas et al. (2019) found that the
addition of KGM in the water phase can significantly increase the creaming stability of
the emulsion containing 0.1% and 0.2% w/v KGM for 14 days due to the increase of viscos-
ity, which induces the enhanced steric hindrance and force of friction between droplets
and the continuous phase. The authors of the study also suggested that the presence of
0.1% and 0.2% (w/v) KGM has significantly increased the curcumin loading compared to
the system without KGM. Moreover, the presence of KGM had no impact on anti-oxidant
activities both in α, α-diphenyl-β-picrylhydrazyl (DPPH) and Ferric Reducing Antioxidant
Power (FRAP) analysis, however, due to the relatively stable curcumin loading capacity,
the anti-oxidant activities of curcumin emulsion remained stable over 14 days of storage.
The particle size and zeta potential value indicated that the addition of KGM had a sig-
nificant impact on decreasing the droplet size, but had no significant impact on the zeta
potential. Lastly, introducing KGM to the emulsion system contributed to slowing down
the release of curcumin from the emulsion droplet during the small intestine digestion
phase due to the presence of the KGM structure in the water, which interfered with the
hydrolysis of the oil droplet (see Figure 10.2).
The results show that the bioaccessibility of curcumin emulsions containing 0.1% and
0.2% KGM (w/v) was lower than that of emulsions without KGM. The low bioaccessibil-
ity means that there is a small concentration of curcumin present in the mixed micelle
compared to the concentration of curcumin in the raw digesta. Furthermore, the results
indicate that introducing KGM to the emulsion system contributes to a slower release of
curcumin from the emulsion droplet during the small intestine digestion phase. The pre-
vious study by Lu et al. (2018) reported that adding KGM significantly modified the release
of β-carotene from the emulsion droplet after passing through the gastrointestinal tract
(GIT). The release of β-carotene was inferior than that of the emulsion without KGM, and
the release rate lowered with the increasing KGM level. Another study by Mao et al. (2015)
also showed that adding a polysaccharide, maltodextrin, to structure the water phase
led to the significantly different release conditions of the hydrophobic food flavour by
FIGURE 10.2
Bioaccessibility of curcumin emulsion during storage. (From Hayuningtyas, A. et al., Effect of konjac gluco-
mannan concentration and oil-phase volume fraction on the stability of curcumin-loaded-oil-in-milk sys-
tem, Proceedings of the 21st Food Innovation Asia Conference 2019 (FIAC 2019), 13–15 June 2019, BITEC, Bangkok,
Thailand, 2019.)
modifying the droplet mobility within the continuous phase. The higher viscosity and
a chain-like structure that was observed in the emulsion with the presence of KGM can
interfere with the hydrolysis of the oil phase and the surface layer of protein by a steric
hindrance effect (Lu et al. 2018). Also, KGM is non-degradable in the small intestine, but
degradable by β-mannanase, an enzyme generated by colon bacteria (Zhang et al. 2014).
In fact, KGM is a promising candidate for the development of controlled bioactive delivery
systems in the upper part of the GIT to the colon part. This mechanism may explain why
emulsions with the presence of KGM in this study showed a lesser release of curcumin
from the emulsion droplet. The results indicate that introducing KGM to the water phase
of emulsions makes a controlled release of curcumin from emulsions feasible. This can be
proven by considering the concentration of curcumin in the micelle (Figure 10.3), which
suggests that the addition of a low concentration of KGM potentially controls the loss of
curcumin during emulsion formation and within the GIT. The concentration of curcumin
in the micelle represents the amount of curcumin that is ready to be absorbed in the final
curcumin emulsion after passing through the upper part of the GIT. The concentration
of curcumin in the micelle was higher in 0.1% KGM compared to 0.2% KGM and without
KGM. The result suggests that structuring water phase with KGM can protect the cur-
cumin within the droplet during formulation, as well as in the digestion process, and this
related to the higher loading capacity as described above. Findings in this study indicated
that structuring the water phase with a low concentration of KGM could make it possible
to develop a curcumin in the milk system containing MCT oil with the potential emulsion
stability and controlled release of curcumin from emulsion.
Moreover, the low bioaccessibility was also shown when increasing the oil volume frac-
tion from 20% to 30% (v/v). Ahmed et al. (2012) reported that the bioaccessibility of cur-
cumin increased with the increase of the lipid level due to the increase of the total amount
of mixed micelle that was available to solubilize the curcumin. The result in this study
FIGURE 10.3
The concentration of curcumin in the micelle during storage after passing through the GIT. (From
Hayuningtyas, A. et al., Effect of konjac glucomannan concentration and oil-phase volume fraction on the sta-
bility of curcumin-loaded-oil-in-milk system, Proceedings of the 21st Food Innovation Asia Conference 2019 (FIAC
2019), 13–15 June 2019, BITEC, Bangkok, Thailand, 2019.)
may have been due to a higher lipid concentration since there was a considerable amount
of lipids, which were probably not digested. Therefore, some curcumin may not have
been released from the emulsion droplets. Hence, there are two possible physicochemi-
cal mechanisms that could describe the correlation between curcumin bioaccessibility and
oil concentration. First, bioaccessibility will increase with the increase of lipid concentra-
tion as more mixed micelle will be formed. Second, the bioaccessibility decreases with the
increase of lipid concentration due to a larger proportion of oil remaining non-hydrolysed,
resulting in some curcumin not being released from the emulsion droplet into the sur-
rounding mixed micelle. In this study, it was expected that the non-hydrolysed oil would
be further absorbed in the colon. Since the KGM was to be degraded by a colonic enzyme,
β-mannanase, the access to the oil would have been facilitated resulting in the release of
curcumin from the oil droplet. Jeppesen and Mortensen (1998) reported that there were
two reasons that can explain the MCT absorption in the colon. First, the MCT can possibly
be absorbed because it has the ability to dissolve in water. The absorption of water-soluble
fatty acid in colon would be related to the report that the colon can completely absorb C8:0
and almost all of C10:0 at a physiological pH. Second, MCT can be absorbed due to the
backward effect of the colon on fat absorption in the small intestine.
There is a considerable potential for KGM to be used for medicinal purposes, particu-
larly for treatment of non-communicable diseases. The physicochemical characteristics of
KGM allow development of novel foods and especially foods for consumers with spe-
cial needs that demand tailor-made formulations. In the future, health promoting foods,
thanks to their nutritional properties, will increasingly be consumed to protect humans
from disease. Thus, the new task for the KGM industry is to provide healthy functional
ingredients for functional food manufacture. This topic will be dealt with in Chapter 11.
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11
New Trends in the Konjac Flour Industry
Chaleeda Borompichaichartkul, Desi Sakawulan, Patthasarun

Pruksarojanakul, and Phattanit Tripetch
CONTENTS
11.1 Introduction....................................................................................................................... 265
11.2 Properties of Konjac Flour............................................................................................... 265
11.3 Konjac Flour as a Novel Coating Material for Encapsulation..................................... 267
11.4 Konjac Flour as Fibre Source........................................................................................... 272
References...................................................................................................................................... 273
11.1 Introduction
Konjac flour is well-known in many industries including the food, agro-industry, biotech-
nology, pharmaceutical, and chemical industry. This is due to the properties of konjac flour
that allow its applications in a wide range of industries. The main applications are: dietary
fibre, thickening agent, gelling agent, water absorption agent, high molecular weight poly-
mer, film-forming material, and stabiliser. In the manufacturing of a food product, konjac
flour can be used to modify its texture, alter its viscosity, make it more solid-like or gel-like,
vary its flexibility, change its organoleptic properties, such as mouth feel, softer, or harder,
and also make it not thin film (e.g., edible coating). Moreover, nowadays, a consumer is
looking for healthy food products which are green, clean or composed of natural ingredi-
ents, easy to prepare, and have a long shelf life. The growing demand among consumers
for healthy products makes researches and industries focus intensively on health benefits.
Therefore, the new trend of using konjac flour in the industry is no longer limited to physi-
cal characteristics or textural properties, but includes more functional properties either
through new inventions or innovations in processing and product development that lead
to health benefits. Hence, this chapter will provide information about the potential appli-
cations of konjac flour in the agro-industry, food, and biotechnology industries, especially
for health and functional products.
11.2 Properties of Konjac Flour

Konjac flour consists of konjac glucomannan (KGM), which is a natural polysaccharide
isolated from tubers of Amorphophallus spp. plants found in Asian countries, such as
China, Taiwan, Japan, and Thailand (Tatirat and Charoenrein 2011). KGM is widely used
265
in the food, pharmaceutical, chemical, and biotechnology industries (Zhang et al. 2005).
The physicochemical properties of KGM are described in Chapter 7. Moreover, KGM has
desirable properties, such as good film-formation, high water solubility, edibility, and bio-
degradability (Tobin et al. 2012). It has a tendency to create a fine dense network upon
drying (Zhang et al. 2005).
KGM powder itself has different characteristics from other powders that are made from
plants. KGM powder is a polymer that is soluble in water. However, it needs a special
technique to dissolve the KGM powder. A crucial factor in the process of the solubilisa-
tion of KGM powder is the fact that it has to be stirred all the time until the KGM powder
is completely dissolved. Otherwise, should it not be stirred all the time, it would become
solid. Furthermore, the KGM powder solution also exhibits different characteristics than
the other polysaccharides:
• The stickiness and the expansion of the KGM powder enable the water absorption
• When it is used in a solution, the particles will absorb and expand into a high vis-
cosity solution pseudoplastic fluid. It can expand 27–40 times its usual size
• The higher the viscosity of the KGM powder, the higher the expansion rate will be
The ability to absorb water by KGM powder will depend on time and temperature.
If the temperature is rising, it will also directly affect the absorption process and
cause the viscosity of the solution to change rapidly. The study from Tye (1991) showed
that the temperature of 50°C is the most suitable for glucomannan expansion. Moreover,
the increase of the viscosity in glucomannan also affects its ability to expand. It can
be argued that the viscosity of konjac powder will depend on the composition of the
KGM as a raw material and the manufacturing process during the extraction of the KGM
powder. Moreover, the gel formation from the KGM powder produced when the solu-
tion is heated under alkaline conditions is an irreversible process. The alkaline solution
will produce a deacetylation reaction, when it reacts with the acetyl group in KGM. As
a result, a gel will be produced due to the formation of a network of hydrogen bonds.
The study also mentions that the lowest concentration of KGM required to form an alka-
line gel is estimated at 0.5%. The alkaline solution substances that are widely chosen
include potassium carbonate, calcium carbonate, and calcium hydroxide. Furthermore,
Tye (1991) explained that the KGM powder can also be used with gum or other powders.
For example, when it is used with kappa carrageenan, the gel itself can become ther-
moreversible. Using the KGM powder with other substances also leads to new types of
gels that can be used in various circumstances, e.g., the creation of a stabiliser gel. More
details on the gelling properties of KGM in combination with other polysaccharides can
be found in Chapter 7.
The properties described above are well-known characteristics of KGM that can be
applied in many areas depending on the product and purpose of use. So far, the main
health benefits of KGM are associated with its nutritional characteristic as a ‘dietary fibre’.
However, the growing trends towards innovation aimed at developing a healthy diet are
focused on a number of new applications. They involve KGM as a coating material form-
ing a viscous solution, gel, or film that is utilised to encapsulate the bioactive compounds
from plants or animals.
New Trends in the Konjac Flour Industry 267
11.3 Konjac Flour as a Novel Coating Material for Encapsulation

Coating materials or wall materials of microencapsulation can be selected from a natural
or synthetic material depending on the core material and desired characteristics in the
final product. In general, carbohydrates and gums are used as coating materials for micro-
encapsulation by spray drying since they show low viscosity, high solid content, and good
solubility (Silva et al. 2014). KGM as a natural polysaccharide can be used as a wall material
substance (Table 11.1). In microencapsulation, using KGM as the coating material has been
reported as a successful method to protect volatile compounds and increase the encapsu-
lation yield (Yang et al. 2009), preserve anti-microbial compounds (Adamiec et al. 2012),
protect and control the release of andrographolide (Wattanaprasert et al. 2017), maintain
and increase anti-oxidant activity of coffee extract (Sakawulan et al. 2017), and preserve
probiotics (Yanprapasiri et al. 2018).
As KGM is viewed as a suitable material for the encapsulation of bioactive compounds
using the spray drying process for its abovementioned characteristics, the use of KGM
is limited by the fact that it has a very high apparent viscosity when dissolved in water,
even at low concentrations (>2% (w/w)). Therefore, its viscosity needs to be reduced to
an appropriate level (<100 mPa·s) before using it in the spray drying and encapsulation
process.
Several studies have considered more in-depth the application of KGM hydrolysate
(KGMH) as a coating agent in microencapsulation by spray drying. The appropriateness
of KGM as a wall material is restricted by its physical properties, particularly, the high
apparent viscosity at low solid content. To be applicable in spray drying microencap-
sulation, KGM should undergo a hydrolysis reaction to get a lower viscosity appro-
priate for a nozzle spray dryer. A hydrolysis reaction can be done either by acid or
TABLE 11.1
Utilisation of KGM and Other Wall Materials in Spray Drying Microencapsulation
Spray Drying Encapsulation
Core Material Wall Material Condition Efficiency (%) References
Sweet orange oil flavour KGM, gum arabic, Inlet temperature: 60–81 Yang et al. (2009)
and starch sodium 160°C
octenyl succinate
Kaffir lime oil (Anti- KGM and KGM+ Inlet temperature: 56.86 Adamiec et al.
microbial agent) gum arabic 160°C–200°C (2012)
Andrographolide KGM, KGM+ Inlet temperature: 10.82–46.63 Wattanaprasert
β-cyclodextrin, 170°C, outlet et al. (2017)
KGM+ temperature: 85°C
α-cyclodexrtin
Coffee extract KGM, maltodextrin Inlet temperature: N/A Sakawulan et al.
160°C–180°C (2017)
Probiotics KGM, maltodextrin, Inlet air 90 Yanprapasiri
trehalose temperature: 170°C et al. (2018)
enzymatic reaction. The acid hydrolysis of the polysaccharides is the simplest method
to convert KGM to be KGM hydrolysate. However, the reaction is time consuming, and
the high acid content in the residue at the end of the reaction hinders its utilisation in
the food industry. For instance, Huang et al. (2010) used sulfuric acid (3.2 M) at 40°C for
7 days and observed the particle size (nanocrystal) after the hydrolysed KGM was dried
by a freeze dryer. Moreover, Wang et al. (2015) also employed 2.8 M sulfuric acid at
40°C for 7 days and investigated the size of the KGM microcrystal. Meanwhile, Tanaka
et al. (2013) used 1 M of sulfuric acid in a boiling water bath for 5 hours with gentle
stirring every 1 hour to get fully hydrolysed KGM solutions. On the other hand, the
specific work of the enzyme shows that enzymatic hydrolysis can be an excellent way
to get the targeted and controlled hydrolysis of polysaccharides. When compared to
acid hydrolysis, o ligosaccharides from the enzymatical hydrolysis of KGM should be
used, as the KGM hydrolysate stimulates the growth of lactobacilli and bifidobacteria
(Al‐Ghazzewi et al. 2007).
There is a study on using β-mannanase to hydrolyse the (1, 4)-β-D-mannopyranosyl
linkages of glucomannans under the condition of 50°C, pH 5.5 for 24 hours. The enzyme
hydrolyses more than 90% of polysaccharides into oligosaccharides and some monosac-
charides (He et al. 2001). Enzyme modification techniques can be used for the modification
of the viscosity of the KGM solution. A number of techniques were devised to decrease
the viscosity of the polysaccharide solution by enzyme hydrolysis, such as using man-
nanase in the processing of plant fibres. Enzymatic hydrolysis leads to the production of
the oligosaccharides in KGM solution, which is used as a raw material in the food industry
(Table 11.2) (Al-Ghazzewi and Tester 2012; Chen et al. 2013).
Moreover, for applications in the food industry, it is more appropriate to use food-grade
enzymes to hydrolyse the KGM than to use concentrated acid. The KGM structure consists
of four different β-D-(1→4) linkages, which are mannose to mannose, mannose to glucose,
glucose to mannose, and glucose to glucose. Therefore, the macromolecule backbone of
KGM is contained in D-mannopyranosyl residue and D-glucopyranosyl residue. The link-
ages in the KGM backbone can be hydrolysed by mannanase, endoglucanase or cellulose,
and pectinase. However, mannanase is the most common enzyme to depolymerise the
KGM due to its high productivity compared to other enzymes. The predominant mono-
mers of KGM are mannose and glucose at a ratio of 1.6:1, while mannanase breaks the
main chain of β-D-1,4-mannopyranosyl linkages. In consequence, mannanase has a higher
productivity than other enzymes for the same timeframe of hydrolysis. On the other
hand, endoglucanases belong to the cellulose-degrading enzymes and cut the linkage of
β-D-1,4-linkages in a cellulose chain. The key targets of a cellulose-degrading enzyme are
TABLE 11.2
Characteristics of the Enzyme Hydrolyses of Substrate Solution
Enzyme Target of Enzyme References
Mannanase 1,4-β-linked mannose-residues Al-Ghazzewi and Tester (2012)

Pectinase 1,4-α-D-galacturonan Rexová-Benková et al. (1983)
Cellulase 1,4-β-linked glucose-residues Al-Ghazzewi and Tester (2012)
the linkages between glucose to mannose and glucose to glucose. Moreover, pectinase
is in the big group of enzymes that break the chain of glycosidic linkage, especially at 1,
4-α-galactosiduronic linkages. In addition, a report from Cescutti et al. (2002) stated that
the presence of few short side chains may contain galactose residues, which are the target
of pectinase enzyme.
Therefore, the most important limitation for using KGM as a coating material, the high
viscosity of KGM, needs to be reduced, and the best and safest way is to produce enzy-
matic hydrolysate. The development of coating material from KGM to encapsulate or pro-
tect bioactive compounds includes kaffir lime oil-anti-microbial compounds (Adamiec
et al. 2012), andrographolide (Wattanaprasert et al. 2017), coffee extracts (Sakawulan et al.
2017), and probiotics (Yanprapasiri et al. 2018). The condition of drying is also important,
but it often depends on the raw material or bioactive compounds. The range of inlet and
outlet air temperature needs to be tailored to the specific product. As for anti-oxidant
properties of some bioactive compounds from the plant extract, they can be degraded or
formed during the high temperature spray drying. Therefore, the optimisation process for
spray drying conditions as well as the ratio of mixture core to wall material are important
and should draw the attention of the researcher. KGM can be used in combination with
other coating materials to synergise the protective effect, for example, KGM and gum ara-
bic (Adamiec et al. 2012), KGM and maltodextrin (Wattanaprasert et al. 2017; Sakawulan
et al. 2017), and KGM, maltodextrin, and trehalose (Yanprapasiri et al. 2018). It would be
interesting to understand the interaction of KGM with other coating materials that would
decrease or enhance the stability of materials, as well as control release. In their study,
Hayuningtyas et al. (2019) found that to stabilise curcumin, which is dissolved in lipids in
the milk system, the addition of KGM (<1%) can make stabilisation possible for this mix-
ture up to 14 days without separation due to the structuring of the water phase with a low
concentration of KGM.
Besides using KGM as a coating material to make a powder or liquid product, there is
another approach that could be applicable for healthy food products corresponding to the
needs of the consumer. An example of such an application could be a functional edible
film that is light, environmentally friendly, safe, and is able to encapsulate and conve-
niently release the bioactive compounds.
KGM is a polysaccharide, which is popular as a raw material for edible films
(Table 11.3). It can form an edible film which is tough, stable, and transparent. In addi-
tion, when mixing other polymers with KGM, they may enhance its mechanical
properties (Mikkonen 2009; Zhang et al. 2005). Other polymers are pullulan, gellan,
polyacrylamide, gelatin (Xiao et al. 2001b), carboxymethyl cellulose (Xiao et al. 2001c,
2002), chitosan (Tripetch et al. 2016; Xiao et al. 2000a), sodium alginate (Xiao et al. 2000b,
2002), polyvinyl pyrrolidone (Xiao et al. 2001a), cellulose, as well as whey protein iso-
late (Leuangsukrerk et al. 2014), etc. Plasticisers, which are often used with KGM edible
film, are glycerol and sorbitol. However, without a plasticiser, KGM can form edible
film alone. The deacetylation of the KGM solution with alkaline solution also increases
the strength of the edible film.
Applications of konjac glucomannan edible film in entrapping bioactive compounds
to preserve or extend shelf life of perishable products include KGM film incorpo-
rated with galangal extract to be used with mangos (Rattananin 2011), KGM film
incorporated with basil oil to be used in leafy salads (Saeheng 2016), and KGM film
TABLE 11.3
Example of KGM Use for Film Forming
Formulation
Polymers Plasticiser Bioactive Compounds References
KGM+gelatin – Xiao et al. (2001b)

KGM+chitosan Glycerol – Xiao et al. (2000a)
KGM+chitosan+soy protein Jia et al. (2009)
isolate
KGM+chitosan Glycerol 5-ALA Tripetch et al. (2016)
KGM+sodium alginate – Xiao et al. (2000b)
KGM Glycerol Basil oil Saeheng et al. (2016)
KGM Glycerol Galangal extract Rattananin (2011)
KGM+carboxymethycellulose – Xiao et al. (2001c)
KGM+gellan – Xu et al. (2007)
incorporated with 5-aminolevulinic acid (5-ALA) to be used in stimulating seed ger-

mination (Tripetch et al. 2016). Rattananin (2011) found that galangal extract entrapped
in KGM film could be released slowly and prevent anthracnose disease in mangoes
destined for export for up to 30 days of storage at 13°C. Colletotrichum gloeosporioides is
the cause of anthracnose disease on mango cv. Nam Dok Mai after harvesting. It was
found that konjac film solution incorporated with A. galanga and B. pandurata extracts
required 10,000 µg/mL as a minimum concentration to inhibit C. gloeosporioides. Konjac
film incorporated with A. galanga can retard anthracnose disease when compared with
a control during 30 days of storage. Meanwhile, Saeheng et al. (2016) found that KGM
film incorporated with basil oil at 4%–6% v/v can inhibit Escherichia coli with a clear
zone size of 9.5–10.1 mm. Tripetch et al. (2016) also found that KGM film can entrap
5-ALA in its matrix as the proposed mechanism (Figure 11.1), as well as stabilise it
during storage. 5-ALA is a known plant regulator and growth promoter. It is a very
sensitive and highly unstable compound and easy to deteriorate. Films from KGM,
KGM-treated with alkali solution (KGOH), chitosan (CHI), as well as blends between
KGOH and CHI were fabricated for 5-ALA entrapment. It was found that the efficiency
of KGM film, KGOH film, and CHI film for 5-ALA entrapment was 55.7% ± 0.73%,
58.3% ± 0.36%, and 60.3% ± 0.18%, respectively. A 25:75 (%w/w) blended film (KGOH/
CHI) showed the highest entrapment efficiency of 5-ALA (65.9% ± 0.37%) vs other films.
The possible mechanism for entrapment of 5-ALA in blended film was postulated
under two mechanisms. A secondary amide that leads to the interaction between the
amino group of CHI and carboxyl group of 5-ALA is proposed as the first mechanism.
The fact that the 5-ALA molecule was entrapped within the complexity of the KGOH
structure is proposed as the second mechanism. Therefore, stabilising 5-ALA in a film
may be an alternative way to use and preserve 5-ALA for further applications.
The abovementioned research results demonstrate that KGM can be used in a film form
to entrap and release bioactive compounds to the targets for preservation or shelf life
extension of various products. In addition to that, KGM is a natural dietary fibre that can
have important functions in the human gut system.
FIGURE 11.1
The proposed mechanism of 5-ALA entrapment in KGM and chitosan film.
11.4 Konjac Flour as Fibre Source

Functional fibres are non-digestible carbohydrates that have a beneficial effect on the
human body. According to chemical characteristics, dietary fibres are classified as soluble
and insoluble fibres. Insoluble fibres are mainly plant cell wall components, such as cel-
lulose, lignin, and hemicellulose, while soluble fibre consists of non-cellulosic polysac-
charides, including pectin, gums, and mucilage (Kalyani Nair et al. 2010). The positive
effects of dietary fibre can be divided into three mechanisms, including bulking, viscosity,
and fermentation. Generally, insoluble fibre exhibits a bulking effect, which can increase
stool mass, alleviate constipation, and maintain bowel movements. Cellulose and lignin,
with their particle formation and water holding ability, are effectively increasing faecal
bulk, reducing intestinal transit time, and preventing and relieving constipation (Ho et al.
2012). Meanwhile, soluble dietary fibre, which is fermentable in the human intestine, may
increase stool bulk by promoting the growth of gut microflora and producing gas and
short-chain fatty acids. Previous studies claim that soluble fibres lower cholesterol, while
insoluble fibres are contributing to stool texture. However, this conclusion is inconsistent,
since not all soluble dietary fibres have significant effects on lowering cholesterol levels,
such as wheat bran and resistant starch, which are classified as soluble dietary fibres, but
these fibres did not appear to affect serum lipids (Jenkins et al. 1998; Slavin 2005). Another
terminology for fibre classification is the viscosity of fibres. A meta-analysis evaluated
the effect of pectin, oat bran, guar gum, and psyllium (2–10 g/day) on blood lipids and
showed significant decreases in total and low-density lipoprotein cholesterol concentra-
tion (Brown et al. 1999).
Glucomannan is classified as a soluble, fermentable, and highly viscous dietary fibre
(Keithley and Swanson 2005; Dikeman and Fahey 2006). The review of glucomannan and
its role in weight reduction was conducted by Keithley and Swanson (2005), and it is
the authors’ conclusion that at the doses of a 2–4 g/day, glucomannan was effective in
inducing a significant weight loss in overweight and obese persons. The glucomannan
molecule in konjac powder was able to promote satiety by delaying gastric emptying and
slowed small-bowel transit time. Glucomannan as a soluble fibre also supports faecal
energy loss by reducing fat and protein absorption (Baer et al. 1997), as well as inhibits
carbohydrate absorption and improves glycaemic parameters (Jenkins et al. 1994). It was
observed that konjac glucomannan also acted as a prebiotic, which promotes human
intestinal Clostridium and produces glucomanno-oligosaccharides followed by short-
chain fatty acid fermentation by gut microbiota (Nakajima et al. 2002). More recently,
Chen et al. (2008) conducted clinical studies with seven constipated Chinese women by
feeding them with 4.5 g/day of konjac glucomannan, which resulted in a significant rise
in weekly defecation frequency and improved the bowel movement. Moreover, they also
found that lactobacilli and short-chain fatty acids were increased resulting in lowering
faecal pH. Hydrolysed konjac glucomannan, which was obtained from a hydrolysis reac-
tion of konjac powder, has exhibited similar benefits as prebiotics. Al‐Ghazzewi et al.
(2007) found that hydrolysed konjac glucomannan was able to promote the growth of
probiotic strains of lactobacilli and bifidobacteria in a single culture. In line with this
study, Connolly et al. (2010) noted that the ability of hydrolysed konjac glucomannan to
promote digestive health was comparable to inulin. In addition, in another study, it has
been reported that the activity of Lactobacillus casei was enhanced with KGM as a prebiotic
(Pruksarojanakul et al. 2017).
The prebiotic activity score is an interesting in vitro method to analyse the prebiotic prop-
erty of the tested saccharide because the data can present the activity of a prebiotic by com-
paring the growth of a probiotic with an enteric mixture. Moreover, Harindhanavudhi
et al. (2019) found that KGM and KGMH have a positive prebiotic activity score with L. casei.
KGMH is a mixture of oligosaccharides produced from the enzymatic hydrolysis of KGM
powder. It can be used as an effective biopolymer coating agent for bioactive materials.
The study of Harindhanavudhi et al. (2019) aimed at studying the effect of KGMH from
15% to 25% (w/w) KGM concentrations on the prebiotic activity score and growth rate of
Lactobacillus casei-01 in goat milk before using it as a coating material for L. casei-01 via spray
drying in a goat milk system. The KGM was hydrolysed by a β-mannanase enzyme and
manno-oligosaccharides were the product of the hydrolysis. The target product from the
hydrolysis was manno-oligosaccharides with 4°–6° of p olymerisation. The results from the
thin-layer chromatography technique demonstrated that by increasing the concentration of
KGM from 15% to 25%(w/w), this increased the concentration of m anno-oligosaccharides,
as shown by the intensity of the spot. The average growth of L. casei-01 was enhanced by
3.71 log CFU/mL (±0.53) when using KGMH as the prebiotic in De Man Rogosa and Sharpe
(MRS) broth. When comparing in vitro prebiotic activity scores between KGMH and com-
mercial prebiotics (inulin and fructo-oligosaccharide), it was found that KGMH from 25%
KGM exhibited a 0.35 ± 0.01 score, which was higher than those of fructo-oligosaccharide
(0.12 ± 0.02) and inulin (−0.02 ± 0.00). Furthermore, KGMH from 25% KGM promoted
L. casei-01 growth in a goat milk system more than KGMH from KGM of 15% and 20%.
These results indicated that KGMH produced from β-mannanase hydrolysis of 25% KGM
could have a potential to be applied in a synbiotic system with L. casei-01. The outcome of
this study shows that KGM and KGMH provide a significant amount of functional proper-
ties to the human body.
Regardless of a direct function in body well-being, KGMH could also be a benefit in food
drying as a drying aid and bioactive encapsulant. Synbiotic is a system where a probiotic
and its prebiotic are present together. Considering KGM or KGMH as a prebiotic of L. casei,
the synbiotic system of L. casei can be built and used as a functional ingredient in food
products.
The new trends in the konjac flour industry are associated with the needs or demands of
consumers that require a healthy product. In addition, the manufacturing process of that
particular product should be green and have zero waste. KGM flour exhibits a number of
advantages to health by adding functional properties to a food product, besides its physi-
cal function in the food product formulation. The examples of the studies mentioned in
this chapter could provide the food manufacturers with information about new trends
in product development and indicate the way to use konjac flour to produce healthy and
functional food products.
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12
Concluding Remarks
KGM is an attractive dietary fibre. It acts as a neutral hydrocolloid with significant health
functions. It has been part of the human diet in China and Japan for nearly 2000 years.
Initially, the main species was Amorphophallus konjac from which the common English
name ‘konjac’ is derived. Nowadays, it has wide applications in the food industry as a
gelling agent, stabilizer, and emulsifier, and is used for edible coatings for the preservation
of fruit and vegetables. Other uses are in the pharmaceutical industry for microencapsula-
tion of active compounds that are to be released ‘on demand’ for treatment of various dis-
eases. There are also applications in the body care and tissue culture field, just to mention
a few of them.
In spite of the many applications, not enough is known about the raw material, i.e., vari-
ous species of the genus Amorphophallus in industrial production, processing, and various
applications of KGM-derived products. There are various publications dedicated to konjac
in the major producing countries like China and Japan in the languages of these countries.
However, even though it is a part of various scientific papers on very specialized topics,
there are so far only a few publications in the English language.
In order to fill in this gap, this book covers a wide range of aspects related to the produc-
tion of KGM. The first of these is the botanical background of the genus Amorphophallus, its
ecology, physiology, and field production. The next major aspect is the postharvest, pro-
cessing, and physicochemical properties of KGM. This is followed by the description on
the latest developments in the drying technology that are the key in the improvement of
the quality of KGM. The next section includes reports on the status of the konjac industry
in the major producing countries. Another major section refers to the applications of KGM
in food and medicine. These areas are the focal point of the research in several countries,
among them China, Thailand, and Indonesia, the latter two increasing considerably their
production of KGM and developing their local manufacturing of the derived products.
Finally, there is a section on the latest trends in the industry that illustrate the application
of the results of the current research done by scientific institutions, and the industry also
doing its own R&D.
The authors of this book very much hope that their work will promote knowledge about
KGM, its source, and its multiple applications.
277
Index
Note: Page numbers in italic and bold refer to figures and tables, respectively.
A Amorphophallus coaetaneus, 25, 26, 27, 28

Amorphophallus corrugatus, 28, 29, 30, 30–31
accumulated temperature, 134
Amorphophallus dunnii, 31, 31, 32, 33, 33
acetate groups, 190
Amorphophallus kachinensis, 33–34, 34, 35, 36
acetyl groups, 197
Amorphophallus kiusianus, 36, 36, 37, 38, 38
acid hydrolysis, 268
Amorphophallus konjac, 1, 38–40, 39, 40, 122, 189,
ADP (Adenosine diphosphate)-glucose, 105–107
238–239
pyrophosphorylase, 110–111
improving mechanization level, 241
adsorption isotherm, KGM, 194
new varieties, breeding, 240
African clade (AFR-subgenus Afrophallus), 7
planting areas, development, 239
agricultural industrialization, 231
promoting professional production, 239–240
agroforestry-based production system,
promoting standardized planting, 240–241
Indonesia
Qingjiang, 123
cultivation, 154
Wanyuan, 122
environment, 152–153
Amorphophallus krausei, 40–41, 41, 42, 43, 43
harvesting of tubers, 154–155
Amorphophallus macrorhizus, 43–44, 44, 45, 46
replanting, 155–156
Amorphophallus muelleri, 46–47, 47, 48, 49, 50,
socio-economic aspects, 156
78, 246
sustainability, 153
species and varieties in China, 124–125
Akagi-ohdama (Nonglin 2), 127–128
Amorphophallus paeoniifolius, 50–51, 51, 52, 53, 54
albinism, 84
Amorphophallus pygmaeus, 54, 55, 56, 56
Amorphophallus, 1; see also specific species
Amorphophallus tenuistylis, 56–57, 57, 58, 59, 59
evolutionary and genetic aspects, 6–9
Amorphophallus thaiensis, 59–60, 60, 61, 62
fruits, 14
Amorphophallus tonkinensis, 62, 62, 63, 64
glucomannan content in, 77–80
Amorphophallus variabilis, 64, 64, 65, 66, 66–67,
glucomannan producing species, 14–77
245–246
growth cycles, 77
Amorphophallus xiei, 67, 67, 68, 69, 70
hybridization, 80–94
Amorphophallus yuloensis, 70, 70, 71, 72, 72
inflorescence, 12–13
Amorphophallus yunnanensis, 72–73, 73, 74, 75,
leaf blade, 12
76, 77
leaflets, 12
amplified fragment length polymorphism
morphology, 6
(AFLP) markers, 9
petiole, 12
anti-browning agents, 165–166, 168–169
phylogenetic tree, 8, 79
anti-browning solution, pre-treatment, 180–181
phylogeny, 7
anti-microbial agents, 147–148
spadix in, 13
antisense gene expression inhibition process,
spathe in, 13
111, 111
species/geographic distribution, 245–246
anti-swelling solution, pre-treatment, 180
taxonomy, 9–11
appendix, 13–14
tubers, 11
Araceae, 13
Amorphophallus albus, 14, 15, 16, 16, 78, 121–122
area selection, cultivation, 142–143
Amorphophallus amygdaloides, 16–17, 17, 18, 19, 19
ascorbic acid, 166–167
Amorphophallus asterostigmatus, 19–20, 20, 21, 22
automation process, level, 242
Amorphophallus bulbifer, 22, 23, 24, 25, 78, 108
279
280 Index
B continental Asia I (CA-I-subgenus Metandrium)

clade, 7
belt dryers, 167, 168
continental Asia II (CA-II-subgenus
bioaccessibility, 261, 261–262
Scutandrium) clade, 7
biological/ecological factors, 239
continuous belt dryer, 246
biosynthesis, KGM, 102
coordination/management in konjac industry
carbohydrates in, 102–104
key players in, 232
enzymes role, 105–106
market prices, 237–238
pathway, 106–110
publicizing, 233–234
soluble glycosides in konjac corm, 104–105
scientific research and international
Bittyu variety, 126, 129
exchanges, 234–237
breeding methods
seminars and conferences, 232
biotechnology, 120
corn starch-konjac systems, 200
crossbreeding, 120
counter-flow tunnel dryer, 246
mutations, 119
crop rotation, 145
plant selection and intraspecific
crossbreeding, 120
varieties, 119
crushing/grinding, 181
varieties, 240
CSLA (cellulose synthase-like family A),
Brown, N. E., 10
107–108, 110
browning, konjac corms
cultivation, 120–121
anti-browning agents, 166–167
Amorphophallus konjac in Japan, 238–241
food quality, 163
areas selection for, 142–143
konjac powder, 164
in China, species and varieties,
phenolic compounds, 163–164
121–125
PPO, 164–166, 165
environment selection for, 143
bulbils, replanting, 155, 155–156
field, 145
burr mill, 247
hazards, 138–142
India, 130
intraspecific varieties, 119
C
in Japan, species and varieties, 125–129
calcium, 136 konjac growth cycle, 154
caramelization reaction, 166 pattern and cropping system, 144–146
cellulose synthase-like family A (CSLA), Southeast Asia, 129
107–108, 110 techniques, 142
centrifugal force, 181 variety improvement, 116–120
centrifugal moisture extractor, 215
centrifugal technique, 215–217, 216, 217
D
Chen, J. F., 121
chitosan (CHI), 270, 271 Da Yu Hu Nan, 103
Chua, M., 166 Dayu variety, 125
Chumohua 1, 124 deacetylation reaction, 197
Claudel, C., 7, 11 digital temperature controller, 209
climate, 239 disease prevention, 226
Clostridium, 272 ditching, 148
coating materials, 267 D-mannose-6-phosphate keto-isomerase, 106
colloid mill, 181 Dowex anion exchange resin, 103
combined wet/dry processing, 178–180, 179 dried konjac chips, 246
common fine flour, drying, 181–182 drought damage, 141
common konjac fine flour, 169, 175 dry extraction, 209
common konjac flour, 169 drying techniques, developments, 228–229
complexation, 202 drying technology, advances in
confocal electron scanning microscopy, 201 konjac flour, 211–220
conical ball mill, 247 konjac tubers, 209–211, 211
Index 281
dry processing/process, 167–169 fresh/dry konjac chips, price, 237

grinding/sifting, 175, 176 fructose, 104, 106, 112
konjac corms, 175 fructose-6-phosphate, 107
step description, 183 functional fibres, 272
E G
elite breeding, 132 Galloway, A., 83
establishment, 132 gastrointestinal tract (GIT), 260, 262
local production/commercialization, 132–133 GDP (Guanosine diphosphate)-mannose, 105,
principles, 131–132 107–108
sterilized konjac seed material, 132 gelatinization, 199
Emoyu 1, 125 gelation mechanism, konjac, 191–192, 192, 193
encapsulation process, 202, 259 gelling agent, 191
novel coating material for, 267–271 generally recognised as safe (GRAS), 2
endotherms, 203 genetic resources
Engler, A., 10 collection and collation, 117
enthalpy, 203 innovative development, 118
environmental compliance standards, 229 GIT (gastrointestinal tract), 260, 262
environmental conditions, 138, 143 glucomannans, 1, 101–103, 180, 189, 272; see also
environmental protection, 243 konjac glucomannan (KGM)
enzymatic browning reaction, 164–166 in Amorphophallus species, 77–80
enzymatic degradation, 110 biosynthesis, 104–105, 107, 109
enzymes, 105–106, 111 in idioblast, 102
glucomannan and starch biosynthesis on, 108 molecules, 196
hydrolysis, 268, 268 producing species, 14–77
modification techniques, 268 and starch biosynthesis, 104, 108, 109
ethanol recycling, 182–183, 183 glucose, 104, 112
glucose-6-phosphate, 107
GRAS (generally recognised as safe), 2
F
Grayum, M. H., 11
Ferrer, O. J., 165 grinding/grinder
fertilizing, 149 konjac chips, 176
field cultivation, 145 mill, 181
field management, 149–150 separation, particles, 183
field production, developments sifting, 175, 176
breeding variety, 224–225, 225–226 techniques, 229
disease prevention, 226
planting areas, 224, 225, 228
H
planting industry, guidelines, 228
planting mode, 226 Halsey model, 186
planting standardization, 227, 227 Haruna-kuro (Nonglin 1), 127–129
film forming, KGM, 270 harvesting, 150
Finne, G., 165 hazards, konjac cultivation
5-ALA, 270 damages caused, wind/hail, 141–142
FLint2 phylogeny, 7 diseases, 138–140
fluidised bed dryer, 216 drought damage, 141
fluidization technique, 209, 210, 216 insect attacks, 140
food/medicine, KGM application, 258–262 low and high temperature, 140–141
food technology, 259 meteorological, 140–141
foreign materials, separation, 243 waterlogging damage, 141
formation process, 102 healthcare industry, 232
free water, 184 Henderson model, 186
282 Index
heterosis, 83 in food, 230

Hetterscheid, W. L. A., 10–11, 14 gel-based foods, 230
Huang, 201 konjac flour (KF), 1–2, 110, 256, 265
hybridization dry processing, 210
in Amorphophallus, 80–94 as fibre source, 272–273
technique, 120 gum, 173, 174
hydrogen bonds, 192 novel coating material, 267, 267–270
hygroscopic character, 184 price, 238
properties, 265–266
quality control/storage, 184–187
I
yield, improving, 242
Impaprasert, R., 166 konjac glucomannan (KGM), 1–2, 78, 190, 191,
induction period, 192 209, 265, 277
industrial applications, 219 biosynthesis, 102–110
insoluble fibres, 272 chemical structure, 191
intercropping system, 145–146 CHI film, 271
-derived products, utilization, 248–249
different temperatures, 197
J
enzymatic degradation, 110
Jackson, S., 83 European Union specifications for, 190
Janovitz-Klapp, A. H., 166 film, 269–270, 270
Jinenjo, 239, 240 food, utilization, 256–257, 258
formation process, 102
gelation mechanism, 191–192, 192
K
gelling agent in food industry, 230
kerupuk puli, 249 gum, different concentrations, 196
Kesatuan Pengelola Hutan (KPH) Saradan, legislation in Europe, 190, 190
152–153, 154, 245 medicine, utilization, 257–258, 258
KF, see konjac flour (KF) -milk protein stabilization mechanism,
KF, dry extraction 259–262
centrifugal technique, 215–217, 216, 217 particles characteristics, 102
disadvantage, 211 physical properties, 193–199
microwave vacuum drying, 217–220 physicochemical properties, 255–256
multistage drying, 212–215, 213 scanning electron micrographs, 220
wet processing of, 212 species with, 78
KGM, see konjac glucomannan (KGM) structure, 190–191
KGM-treated with alkali solution (KGOH), 270 synthesis, regulation, 110–112
Koch, K., 40 utilization, 252
konjac, processing methods, 246 konjac growth, 104
machinery for, 246–247 konjac industry
mechanical drying, konjac chips, 246 in China, 224–238
sun drying, konjac chips, 246 in Indonesia, 245–249
by wet process, 247–248 in Japan, 238–245
konjac corms, 211 structure, 250–252, 251
browning control, 163–167 in Thailand, 250–252
drying/storage, dried chips, 167–169 konjac noodles, 248
harvesting and storage, 161 konjac planting areas, development, 239
primary processing, 162, 162–163 konjac processing, principles,
processing plant, delivery to, 162 173–174, 174
soluble glycosides in, 104–105 konjac produced/consumed, Thailand, 250
konjac derived products, development konjac products industry, Japan
domestic/international markets, automation degree, 244
synchronized expansion, 230–231 creating awareness, 244
Index 283
new products development, 245 native variety, 125–126

overview, 243–244 natural growth method, 144–145
konjac tofu, 248 natural polysaccharide, 267
KPH (Kesatuan Pengelola Hutan) Saradan, new products development, 245
152–153, 154, 245 ‘Nightstick,’ 40
nitrogen, 136
non-reversibility property, 199
L
non-scrapping washing method, 163
lactic acid, 249 non-sessile stigma, 7
‘Leo Song,’ 40 non-sulphite anti-browning agents, 166
Li, H., 14 normalised heating scans, 203
light intensity, 134–135 nucleotide sugars, 105
Liu, P. Y., 121
O
M
offset tubers, 121–123
macromolecules, 179 corms and, 131–132
Madero, C. F., 165 rapid growth technique for, 132
Maillard reaction, 164 reproduction by, 146
mannose, 103, 106 okayama-ken variety, 126
market prices, 237–238 oligosaccharides, 110
MCT (medium chain triglyceride), 260 orange konjac jellies, 256
mechanization level, 241 organic fertilizer, 137
medium chain triglyceride (MCT), 260 organic solution, 177
Mekkerdchoo, O., 78
microencapsulation, 259, 267; see also
P
encapsulation process
microsatellite marker, 7, 9 packaging, 183
microwave energy, 219 Parry, J., 2
microwave-vacuum drying techniques, pest control, 149–150
217–220 phosphoglucomutase, 106
mineral nutrition, 136–137 phosphomannose isomerase, 106
moisture absorption curve, 186 phosphorus, 136
moisture conditions, 135 photosynthesis, 103–105, 107
moisture content, 216 physicochemical mechanisms, 262
moisture sorption isotherms, 185 planting area, 224, 225
molecular docking analysis, 248–249 planting industry, 232
molecular markers, 7, 9 committee, 236
monosaccharides, 104, 110 guidelines, 228
mutation breeding, 121 standards for konjac, 227
Myogi-yutaka (Nonglin 3), 129–130 planting mode, 226
planting standardization, 227, 227
N planting time/density, 148
plot preparations, 148
Nakajima Taniwa, 173 pollen bank, 83
Nanyang variety, 125 pollen fertility, 81–84
national macro-policy, guiding role polyphenol oxidase (PPO), 164–166, 165
agricultural industrialization, 231 polysaccharides, 266, 269
healthcare industry, 232 acid hydrolysis, 268
on konjac-related themes, 232, 233 gelling properties with, 199–203
poverty alleviation, 231 glucomannan, 103, 190
publicizing konjac industry, proteins and, 259
233–234 stabilization, 260
284 Index
porang, 152 Shimahara, H., 166

potassium, 136 Shimizu, N., 166
poverty alleviation, 231 Shiyu variety, 125
powdered food, 184 Siebold, P. von, 40
PPO (polyphenol oxidase), 164–166, 165 sigmoid/S-shape, 186
prebiotic activity, 273 soft rot, 138–139
processing industry, current techniques soil, konjac cultivation, 137, 143–144, 239
drying techniques, developments, 228–229 loosening, 149
environmental compliance standards, 229 physical/chemical characteristics,
konjac production in southeast Asia, 229–230 144, 144
sulphur dioxide content, standards, 229 preparation, 148
processing industry, Japan, 241–242 surface coverage, 149
coordinating role, 243 soluble dietary fibre, 272
environmental protection, 243 sorption isotherms, 184–185
foreign materials, separation, 243 Southeast Asian (SEA-subgenus
konjac flour yield, improving, 242 Amorphophallus) clade, 7
process automation, level, 242 southern blight, 139–140
promoting professional production, 239–240 spadix, 13
promoting standardized planting, 240–241 spathe, 13
proteins stabilization mechanism, 259
adsorption, 260 starch, 103–104
CSLA3, 109–110 starch-konjac mixed systems, 199–201
and polysaccharides, 259 sterilization, seed konjac, 147–148
-stabilized emulsion, 259 sterilized konjac seed material, 132
publicizing konjac industry, 233–234 storage
conditions, 151
curing before, 150–151
Q
insulated warehouse, 151
Qinmo 1, 124 open field protection, overwintering, 152
outdoor piled, 152
physiological changes, 150
R
simple indoor tempering, 151–152
random amplified polymorphic DNA (RAPD) strong sunlight, 141
markers, 7, 9, 78 substrate solution, enzyme hydrolyses, 268
reaction time, determination, 180 sucrose, 103–104
reproduction methods, 146–147 decomposition, 112
rhizomatous offsets, 146 synthase, 105–106
ridging, 148 sulphiting agents, 165
rotary cutting mill, 247 sulphur dioxide content
Ruo Lin Zhong Dao, 102 residues, 214
standards, 229
sulphur fumigation, 229
S
sun drying, konjac chips, 246
Schott, H. W., 9
scientific research/international exchanges
T
konjac industry development, 237
and technological achievements, 234, Tak province, konjac, 252
235–236 Tan, J., 82
screw mill, 247 targeted poverty alleviation, 231
secondary drying process, 212 temperatures, 133–134
seed konjac selection, 147 accumulated, 134
shade crops, 137–138 digital controller, 209
shear thinning phenomenon, 195 KGM, 197
Index 285
test-tube plantlet, 131 water solubility/absorption, 193–194, 194

thermally stable gels, 197–199, 198 weeding, 149
thickening agent, 195–197, 196 wet method, 173
three-dimensional planting mode, 226 wet milling, 180
three-stage breeding system, 131 wet processing, 177, 180
tissue culture technique, 147 anti-browning solution, 180–181
topography, 239 anti-swelling solution, 180
Toronto Zoo, 83 common fine flour, drying, 181–182
trace elements, 136 crushing/grinding, 181
traditional konjac products, 230 ethanol recycling, 182–183, 183
Tripetch, P., 270 fresh corms, raw material, 177
tunnel dryer, 246 KF, raw material, 178
reaction time, determination, 180
washing/dehydration, 181
U
whiteness index value, 214
UDP (Uridine diphosphate)-glucose, wind, 137
105, 107
X
V
X-ray diffraction spectra, 203
vacuum drying method, 182–183
vibrating-fluidized bed dryers, 167, 169
Y
viscosity, konjac flour, 217
yellow konjac, 125
Yumo 1, 122–123
W
Yunyu 1, 123–124
Walker, J. R. L., 165
washing/dehydration, 181
Z
washing unit, konjac corms, 162, 163
water/cryoprotective properties, 203–204 Zhang, D., 164, 177
waterlogging damage, 141 Zhang, S., 80
water-miscible organic solvent, 212 Zhao, J., 166

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Konjac Glucomannan

© 2020 by Taylor & Francis Group, LLC

No claim to original U.S. Government works

Printed on acid-free paper

International Standard Book Number-13: 978-1-138-36717-3 (Hardback)

Library of Congress Cataloging‑in‑Publication Data

Names: Srzednicki, G. (George), editor. | Borompichaichartkul, Chaleeda,

Visit the Taylor & Francis Web site at

and the CRC Press Web site at

A fine obituary has been published in Biotropica 48(1) in 2016.

Max and Amorphophallus maxwellii

2. Botanical Background to Amorphophallus........................................................................ 5

3. Biosynthesis and Decomposition of Konjac Glucomannan....................................... 101

4. Field Production of Konjac................................................................................................ 115

5. Postharvest Technology of Konjac................................................................................... 161

6. Processing of Konjac Flour................................................................................................ 173

7. Physico-Chemical Properties of Konjac Glucomannan.............................................. 189

8. Advances in Drying Technology...................................................................................... 209

9. Konjac Industry in Major Producing Countries........................................................... 223

10. Applications of Konjac Glucomannan in Food and Medicine................................... 255

11. New Trends in the Konjac Flour Industry...................................................................... 265

12. Concluding Remarks.......................................................................................................... 277

Prof. Zhang Shenglin

Dr George Srzednicki completed a PhD in technical sciences

Chaleeda Borompichaichartkul Zhao Jianrong

Patthasarun Pruksarojanakul George Srzednicki

Chaleeda Borompichaichartkul and George Srzednicki

Konjac glucomannan (KGM) is a high-molecular-weight polysaccharide that was originally

Wilbert Hetterscheid, Li Heng, Wang Zhonglang, Orachorn Mekkerdchoo,

2.1 Evolutionary and Genetic Aspects

Engler (1876) reduced Conophallus, Proteinophallus and Brachyspatha to Amorphophallus

2.2.1 Amorphophallus: Genus Description

2.3 Glucomannan-Producing Species

2.3.1 Amorphophallus albus Liu & Wei

Distribution: China: Sichuan, Yunnan.

NO T E :Amorphophallus albus belongs to subgenus Scutandrium. Although its morphology

2.3.2 Amorphophallus amygdaloides Hett. & Sizemore

Distribution: Southwestern Thailand, Kanchanaburi province.

NO T E : Amorphophallus amygdaloides is a member of a small group of species with broad

2.3.3 Amorphophallus asterostigmatus Bogner & Hett.

is sessile or subsessile (lowermost 1.5–3 mm devoid of flowers), slightly shorter to slightly

Distribution: Central Thailand: Lop Buri prov., Saraburi prov.

N O T E : The inflorescence of A. asterostigmatus is most similar to A. longituberosus but

2.3.4 Amorphophallus bulbifer (Roxb.) Blume

NO T E :Amorphophallus bulbifer is a well-known species in circles of plant lovers. It has

2.3.5 Amorphophallus coaetaneus S. Y. Liu & S. J. Wei

2.3.6 Amorphophallus corrugatus N. E. Br.

Distribution: Northern Thailand, N. Myanmar, SW China (SE Yunnan, Guangxi),

forced by ecological circumstances because A. corrugatus is found in the same ecological

2.3.7 Amorphophallus dunnii Tutch

Distribution: China: Guangdong, Guangxi.

2.3.8 Amorphophallus kachinensis Engl. & Gehrm.

NO T E : See under A. corrugatus (Section 2.3.6).

2.3.9 Amorphophallus kiusianus (Makino) Makino

Distribution: China: Anhui, Fujian, Guangdong, Hunan, Jiangxi; Taiwan; southern

2.3.10 Amorphophallus konjac K. Koch

Distribution: China, Yunnan. It is widely cultivated in China and Japan.

2.3.11 Amorphophallus krausei Engl.

either refrigerated or frozen pollen. This demonstrates the usefulness of refrigerated/